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Holden Clark

Fundamentals of
Weed Science
Fundamentals of
Weed Science
Fundamentals of
Weed Science

Edited by
Holden Clark
Fundamentals of Weed Science
Edited by Holden Clark
ISBN: 978-1-9789-2502-1

© 2017 Library Press

Cataloging-in-Publication Data

Fundamentals of weed science / edited by Holden Clark.


p. cm.
Includes bibliographical references and index.
ISBN 978-1-9789-2502-1
1. Weeds. 2. Weeds--Control. 3. Herbicides. I. Clark, Holden.
SB611 .F86 2017
632.5--dc23

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Table of Contents

Preface  VII

Chapter 1 Understanding Weed 1

Chapter 2 Types of Weed 8


a. Crop Weed 8
b. Noxious Weed 8
c. Abrus Precatorius 11
d. Bromus Tectorum 16
e. Cardoon 18
f. Digitaria Ciliaris 21
g. Juncus Effusus 22
h. Hedera Helix 24
i. Kudzu 29
j. Beneficial Weed 35

Chapter 3 Species of Weed: Invasive and Introduced 111


a. Invasive Species 111
b. Introduced Species 124
c. Chrysanthemoides Monilifera 130
d. Heracleum Mantegazzianum 133
e. Impatiens Glandulifera 136
f. Fallopia Japonica 139
g. Pilosella Aurantiaca 145
h. Echium Plantagineum 146
i. Asparagus Asparagoides 147

Chapter 4 Plants Considered to be Weed 150


a. Amaranth 150
b. Toxicodendron Radicans 159
c. Taraxacum 166
d. Plantago Major 172
e. Silybum Marianum 175
f. Trifolium Repens 178
g. Striga 181

Chapter 5 Weed Control: Methods and Techniques 187


a. Weed Control 187
b. Aquatic Weed Harvester 194
c. Soil Steam Sterilization 197
d. Soil Solarization 202
e. Stale Seed Bed 204
f. Drip Irrigation 205
g. Bioherbicide 211

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VI   Contents

h. Bush Regeneration 212


i. Cultivator 215
j. Mulch 220
k. Hoe (Tool) 227
l. Herbicide 232
m. Glyphosate 247

Chapter 6 Laws Related to Weed 264


a. Weeds Act 264
b. International Plant Protection Convention 265
c. Federal Noxious Weed Act 266


Permissions


Index

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Preface  
Weed science is the part of science, which studies the management of weed growth in the fields
of horticulture and agriculture. The subject includes methods of weed removal as well as the
advantages of the growth of beneficial weeds. The aim of this text is to present the subject of
weed science in the most comprehensive and easy to understand manner. It includes topics
which are crucial for the understanding of the subject. The text includes different approaches,
evaluations and methodologies on weed science. The textbook aims to serve as a resource guide
for students and facilitate the study of the discipline.

To facilitate a deeper understanding of the contents of this book a short introduction of every
chapter is written below:

Chapter 1- Any plant that is considered to be undesirable is known as weed. Some of the examples
of unwanted plants in human setting are lawns, parks and gardens. The chapter on weed offers
an insightful focus, keeping in mind the subject matter.

Chapter 2- Abrus precatorius is a perennial climber that climbs around trees or shrubs. It is
usually known for its seeds. Its seeds can be used as beads and in percussion instruments. The
other types of weed plants are Bromus tectorum, cardoon, Digitaria ciliaris, Juncus effusus and
kudzu. The chapter serves as a source to understand all the types of weed.

Chapter 3- Invasive species are species such as plants, fungi or even animals that are not native
to a particular location. Chrysanthemoides monilifera, Heracleum mantegazzianum, Fallopia
japonica, Pilosella aurantiaca and Asparagus asparagoides are some of the examples given in
this chapter. The chapter strategically encompasses and incorporates the main species of weed,
providing a complete understanding.

Chapter 4- The plants considered to be weed are amaranth, Toxicodendron radicans, tataxacum,
Silybum marianum and striga. Amaranth is a short-lived perennial plant; most of the species
of amaranthus are summer annual weeds. The topics discussed in the section are of great
importance to broaden the existing knowledge on weed.

Chapter 5- Controlling or stopping unwanted plants is termed as weed control. It is very


important for agriculture as weed can compete with domesticated plants. The methods and
techniques discussed in this section are aquatic weed harvester, soil steam sterilization, stale
seed bed, drip irrigation, bush regeneration and herbicide. The aspects elucidated in the chapter
are of vital importance, and provides a better understanding of weed control.

Chapter 6- The Weeds Act 1959 is an act that is regarding the control of several injurious weed
species. This act is restricted to the United Kingdom. International Plant Protection Convention
and the Federal Noxious Weed Act of 1974 are some of the laws related to weed control. This
chapter discusses the laws related to weeds in a critical manner providing key analysis to the
subject matter.

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VIII   Preface

I owe the completion of this book to the never-ending support of my family, who supported me
throughout the project.

Editor

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1
Understanding Weed
Any plant that is considered to be undesirable is known as weed. Some of the examples of unwant-
ed plants in human setting are lawns, parks and gardens. The chapter on weed offers an insightful
focus, keeping in mind the subject matter.

A weed is a plant considered undesirable in a particular situation, “a plant in the wrong place”. Ex-
amples commonly are plants unwanted in human-controlled settings, such as farm fields, gardens,
lawns, and parks. Taxonomically, the term “weed” has no botanical significance, because a plant
that is a weed in one context is not a weed when growing in a situation where it is in fact wanted,
and where one species of plant is a valuable crop plant, another species in the same genus might
be a serious weed, such as a wild bramble growing among cultivated loganberries. Many plants
that people widely regard as weeds also are intentionally grown in gardens and other cultivated
settings, in which case they are sometimes called beneficial weeds. The term weed also is applied
to any plant that grows or reproduces aggressively, or is invasive outside its native habitat. More
broadly “weed” occasionally is applied pejoratively to species outside the plant kingdom, species
that can survive in diverse environments and reproduce quickly; in this sense it has even been
applied to humans.

weed: “A herbaceous plant not valued for use or beauty, growing wild and rank, and regarded as
cumbering the ground or hindering the growth of superior vegetation... Applied to a shrub or tree,
especially to a large tree, on account of its abundance in a district... An unprofitable, troublesome,
or noxious growth.”

-- The New shorter Oxford English dictionary on historical principles

Weed control is important in agriculture. Methods include hand cultivation with hoes, powered
cultivation with cultivators, smothering with mulch, lethal wilting with high heat, burning, or
chemical attack with herbicides.

Ecological significance
Certain classes of weeds share adaptations to ruderal environments, that is to say, disturbed en-
vironments where soil or natural vegetative cover has been damaged or frequently gets damaged,
disturbances that give the weeds advantages over desirable crops, pastures, or ornamental plants.
The nature of the habitat and its disturbances will affect or even determine which types of weed
communities become dominant.

Examples of such ruderal or pioneer species include plants that are adapted to naturally occurring
disturbed environments such as dunes and other windswept areas with shifting soils, alluvial flood
plains, river banks and deltas, and areas that are burned repeatedly. Since human agricultural
practices often mimic these natural environments where weedy species have evolved, some weeds
are effectively preadapted to grow and proliferate in human-disturbed areas such as agricultural

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2 Fundamentals of Weed Science

fields, lawns, roadsides, and construction sites. The weedy nature of these species often gives them
an advantage over more desirable crop species because they often grow quickly and reproduce
quickly, they commonly have seeds that persist in the soil seed bank for many years, or they may
have short lifespans with multiple generations in the same growing season. In contrast, peren-
nial weeds often have underground stems that spread under the soil surface or, like ground ivy
(Glechoma hederacea), have creeping stems that root and spread out over the ground.

A dandelion is a common plant all over the world, especially in Europe, Asia, and the Americas. It is a well-known
example of a plant that is considered a weed in some contexts (such as lawns) but not a weed in others
(such as when it is used as a leaf vegetable or herbal medicine).

Some plants become dominant when introduced into new environments because the animals in
their original environment, that compete with them or feed on them are absent; in what is some-
times called the “natural enemies hypothesis”, plants freed from these specialist consumers may
become dominant. An example is Klamath weed, that threatened millions of hectares of prime
grain and grazing land in North America after it was accidentally introduced, but was reduced to
a rare roadside weed within several years after some of its natural enemies were imported during
World War II. In locations where predation and mutually competitive relationships are absent,
weeds have increased resources available for growth and reproduction. The weediness of some
species that are introduced into new environments may be caused by their production of allelo-
pathic chemicals which indigenous plants are not yet adapted to, a scenario sometimes called the
“novel weapons hypothesis”. These chemicals may limit the growth of established plants or the
germination and growth of seeds and seedlings.

Another of the ways in which the ecological role of a plant can make it a weed even if it is in it-
self inoffensive, is if it harbours a pest that is dependent on it for survival; for example, Berberis
species are intermediate hosts for stem rust fungi, so that they promote serious damage to wheat
crops when growing near the fields.

Competition With Cultivated and Endemic Plants


A number of native or non-native plants are unwanted in a specific location for a number of rea-
sons. An important one is that they interfere with food and fiber production in agriculture, where-
in they must be controlled in order to prevent lost or diminished crop yields. Other important
reasons are that they interfere with other cosmetic, decorative, or recreational goals, such as in

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Understanding Weed 3

lawns, landscape architecture, playing fields, and golf courses. Similarly, they can be of concern
for environmental reasons whereby introduced species out-compete for resources or space with
desired endemic plants. For all these reasons; horticulture, both functional and cosmetic, and en-
vironmental, - weeds interfere by:

• competing with the desired plants for the resources that a plant typically needs, namely,
direct sunlight, soil nutrients, water, and (to a lesser extent) space for growth;

• providing hosts and vectors for plant pathogens, giving them greater opportunity to infect
and degrade the quality of the desired plants;

• providing food or shelter for animal pests such as seed-eating birds and Tephritid fruit flies
that otherwise could hardly survive seasonal shortages;

• offering irritation to the skin or digestive tracts of people or animals, either physical irri-
tation via thorns, prickles, or burs, or chemical irritation via natural poisons or irritants in
the weed (for example, the poisons found in Nerium species);

• causing root damage to engineering works such as drains, road surfaces, and foundations,
blocking streams and rivulets.

700 cattle that were killed overnight by a poisonous weed.

In weed ecology some authorities speak of the relationship between “the three Ps”: plant, place,
perception. These have been very variously defined, but the weed traits listed by H.G. Baker are
widely cited.

Weeds have long been a concern, perhaps as long as humans have cultivated plants. They are men-
tioned in various historic texts, such as a Shakespearean sonnet:

“To thy fair flower add the rank smell of weeds: / But why thy odour matcheth not thy show, / The
soil is this, that thou dost common grow.”

and the Bible:

“Cursed is the ground because of you; through painful toil you will eat of it all the days of your life.
It will produce thorns and thistles for you, and you will eat the plants of the field. By the sweat of
your brow you will eat your food until you return to the ground,”

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4 Fundamentals of Weed Science

Benefits of Weed Species


“What would the world be, once bereft, of wet and wildness? Let them be left. O let them be left;
wildness and wet; Long live the weeds and the wilderness yet.”

-- Gerard Manley Hopkins’ poem Inversnaid

While the term “weed” generally has a negative connotation, many plants known as weeds can
have beneficial properties. A number of weeds, such as the dandelion (Taraxacum) and lamb’s
quarter, are edible, and their leaves or roots may be used for food or herbal medicine. Burdock is
common over much of the world, and is sometimes used to make soup and medicine in East Asia.
Some weeds attract beneficial insects, which in turn can protect crops from harmful pests. Weeds
can also prevent pest insects from finding a crop, because their presence disrupts the incidence of
positive cues which pests use to locate their food. Weeds may also act as a “living mulch”, providing
ground cover that reduces moisture loss and prevents erosion. Weeds may also improve soil fertili-
ty; dandelions, for example, bring up nutrients like calcium and nitrogen from deep in the soil with
their tap root, and clover hosts nitrogen-fixing bacteria in its roots, fertilizing the soil directly. The
dandelion is also one of several species which break up hardpan in overly cultivated fields, helping
crops grow deeper root systems. Some garden flowers originated as weeds in cultivated fields and
have been selectively bred for their garden-worthy flowers or foliage. An example of a crop weed
that is grown in gardens is the corncockle, (Agrostemma githago), which was a common weed in
European wheat fields, but is now sometimes grown as a garden plant.

Some people have appreciated weeds for their tenacity, their wildness and even the work and con-
nection to nature they provide. As Christopher Lloyd wrote in The Well-Tempered Garden

“Many gardeners will agree that hand-weeding is not the terrible drudgery that it is often made out
to be. Some people find in it a kind of soothing monotony. It leaves their minds free to develop the
plot for their next novel or to perfect the brilliant repartee with which they should have encoun-
tered a relative’s latest example of unreasonableness.”

Role in Mass Extinctions


A mass extinction is generally caused by some abrupt disruption to the entire planet’s environ-
ment. This results in major changes in habitat worldwide, and most endemic species, specially
adapted to a single habitat, cannot survive in the new habitats. Thus only weedy species survive,
and they dominate the planet in the immediate aftermath. Cockroaches, for example, have sur-
vived several mass extinctions. The current Holocene extinction event, then, could lead to a planet
inhabited entirely by what are known today as weeds. The fossil record indicates that after mass
extinctions, a weed-dominated planet persists for five to ten million years before life re-diversifies.

Dispersal
Many weed species have moved out of their natural geographic ranges and spread around the
world in tandem with human migrations and commerce. Weed seeds are often collected and trans-
ported with crops after the harvesting of grains, so humans are a vector of transport as well as a
producer of the disturbed environments to which weed species are well adapted, resulting in many
weeds having a close association with human activities.

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Understanding Weed 5

Some weed species have been classified as noxious weeds by government authorities because, if
left unchecked, they often compete with native or crop plants or cause harm to livestock. They are
often foreign species accidentally or imprudently imported into a region where there are few nat-
ural controls to limit their population and spread.

Weeds as Adaptable Species


“We’ve got to be one of the most bomb-proof species on the planet.”

paleontologist David Jablonsky

An alternate definition often used by biologists is any species, not just plants, that can quick-
ly adapt to any environment. Some traits of weedy species are the ability to reproduce quickly,
disperse widely, live in a variety of habitats, establish a population in strange places, succeed in
disturbed ecosystems and resist eradication once established. Such species often do well in hu-
man-dominated environments as other species are not able to adapt. Common examples include
the common pigeon, brown rat and the raccoon. Other weedy species have been able to expand
their range without actually living in human environments, as human activity has damaged the
ecosystems of other species. These include the coyote, the white-tailed deer and the brown headed
cowbird.

In response to the idea that humans may face extinction due to environmental degradation, pale-
ontologist David Jablonsky counters by arguing that humans are a weed species. Like other weedy
species, humans are widely dispersed in a wide variety of environments, and are highly unlikely to
go extinct no matter how much damage the environment faces.

Plants Often Considered to Be Weeds

White clover

White clover is considered by some to be a weed in lawns, but in many other situations is a desir-
able source of fodder, honey and soil nitrogen.

A short list of some plants that often are considered to be weeds follows:

• Amaranth – (“pigweed”) annual with copious long-lasting seeds, also a highly edible and
resilient food source

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6 Fundamentals of Weed Science

• Bermuda grass – perennial, spreading by runners, rhizomes and seeds.

• Bindweed

• Broadleaf plantain – perennial, spreads by seeds that persist in the soil for many years

• Burdock – biennial

• Common lambsquarters – annual

• Creeping charlie – perennial, fast-spreading plants with long creeping stems

• Dandelion – perennial, wind-spread, fast-growing, and drought-tolerant

• Goldenrod – perennial

• Japanese knotweed

• Kudzu – perennial

• Leafy spurge – perennial, with underground stems

• Milk thistle – annual or biennial

• Poison ivy – perennial

• Ragweed – annual

• Sorrel – annual

• Striga

• St John’s wort – perennial

• Sumac – woody perennial

• Tree of heaven – woody perennial

• Wild carrot – biennial

• Wood sorrel – perennial

• Yellow nutsedge – perennial

Weed Control
Weeds are plants that some people view as undesirable in a particular place. Throughout the long
human history of horticulture, people have worked to control weeds for many reasons. Weed con-
trol is a highly developed field of knowledge.

Weed control methods vary according to the growth habit of the weeds in questions, as well as the
context. For example, different methods of weed control may be used on a food crop versus a fiber
crop or a golf course, because there is often more concern about health effects of chemicals used
on food crops, because they are ingested.

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Understanding Weed 7

Weeds can be categorized by their life habit. They can generally either be grouped as annuals or
perennials. An annual weed grows from the seeds dropped in the previous growing season. Peren-
nial weeds regrow from previously established roots, dormant stolons, tubers, rhizomes, as well
as the seed.

If chemical control is chosen as a method of weed control, the grouping of weeds as annual or
perennial is important. If chemical control is chosen, then pre-emergent herbicides would reduce
annual weeds whereas perennial weeds would require the use of post-emergent herbicides.

Understanding the habit of weeds is also important for non-chemical methods of weed control,
such as plowing, surface scuffling, promotion of more beneficial cover crops, and prevention of
seed accumulation in fields. For example, amaranth is an edible plant that is considered a weed
by mainstream modern agriculture. It produces copious seeds (up to 1 million per plant) that last
many years, and is an early-emergent fast grower. Those seeking to control amaranth quote the
mantra “This year’s seeds become next year’s weeds!” However, another view of amaranth values
the plant as a resilient food source.

History
It has long been assumed that weeds, in the sense of rapidly evolving plants taking advantage of
human-disturbed environments, evolved in response to the Neolithic agricultural revolution ap-
proximately 12,000 years ago. However, researchers have found evidence of “proto-weeds” behav-
ing in similar ways at Ohalo II, a 23,000-year-old archeological site in Israel.

In Other Media
• In the video game Plants vs Zombies Garden Warfare 2 there’s a plant called Weed

• There is a song by Snoop Dogg called The Next Episode that refers to smoking weed at the
end.

References
• Brown, Lesley (1993). The New shorter Oxford English dictionary on historical principles. Oxford [Eng.]: Clar-
endon. ISBN 0-19-861271-0.

• Hans Lambers; F Stuart Chapin III; Thijs L. Pons (8 October 2008). Plant Physiological Ecology. Springer. pp.
507–. ISBN 978-0-387-78341-3.

• Annecke, D. R., Moran, V. C. (1982). Insects and mites of cultivated plants in South Africa. London: Butter-
worths. ISBN 0-409-08398-4.

• Muhammad Ashraf; Münir Öztürk; Muhammad Sajid Aqeel Ahmad; Ahmet Aksoy (2 June 2012). Crop Pro-
duction for Agricultural Improvement. Springer. pp. 525–. ISBN 978-94-007-4116-4.

• National Geographic (2011). National Geographic Answer Book: 10,001 Fast Facts about Our World. National
Geographic Society. pp. 175–. ISBN 978-1-4262-0892-8.

• House (U S ) Office of the Law Revision (25 April 2008). United States Code, 2006, V. 3, Title 7, Sections 701-
End. Government Printing Office. pp. 1230–. ISBN 978-0-16-079998-3.

• Ainit Snir; et al. (22 July 2015). “The Origin of Cultivation and Proto-Weeds, Long Before Neolithic Farming”.
PLOS ONE. 10: e0131422. doi:10.1371/journal.pone.0131422. Retrieved 23 July 2015.

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2
Types of Weed
Abrus precatorius is a perennial climber that climbs around trees or shrubs. It is usually known for
its seeds. Its seeds can be used as beads and in percussion instruments. The other types of weed
plants are Bromus tectorum, cardoon, Digitaria ciliaris, Juncus effusus and kudzu. The chapter
serves as a source to understand all the types of weed.

Crop Weed
Crop weeds are weeds that grow amongst crops.

Types of Crop Weed


Examples of crop weeds include chickweed, barnyard grass, dandelion, striga and fallopia japon-
ica.

Effects
Despite the potential for some crop weeds to be used as a food source, many can also prove harm-
ful to crops, both directly and indirectly. Crop weeds can inhibit the growth of crops, contaminate
harvested crops and often spread rapidly. They can also host crop pests such as aphids, fungal rots
and viruses. Cost increases and yield losses occur as a result. Striga, one of the main cereal crop
weeds in Sub-Saharan Africa, commonly causes yield losses of 40–100% and accounts for around
$7 billion in losses annually. Around 100 million hectares of land in Sub-Saharan Africa are affect-
ed by striga. Barnyard grass has been identified as a culprit in global rice yield losses and certain
species have been known to mimic rice.

Noxious Weed
A noxious weed, harmful weed or injurious weed is a weed that has been designated by an agri-
cultural authority as one that is injurious to agricultural or horticultural crops, natural habitats or
ecosystems, or humans or livestock. Most noxious weeds have been introduced into an ecosystem
by ignorance, mismanagement, or accident. Some noxious weeds are native. Typically they are
plants that grow aggressively, multiply quickly without natural controls (native herbivores, soil
chemistry, etc.), and display adverse effects through contact or ingestion. Noxious weeds are a
large problem in many parts of the world, greatly affecting areas of agriculture, forest manage-
ment, nature reserves, parks and other open space.

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Types of Weed 9

These weeds are typically agricultural pests, though many also have impacts on natural areas.
Many noxious weeds have come to new regions and countries through contaminated shipments of
feed and crop seeds or intentional introductions such as ornamental plants for horticultural use.

Invasive weed, mature Scotch Thistle in Australia (person for scale)

Types
There are types of noxious weeds that are harmful or poisonous to humans, domesticated grazing
animals, and wildlife. Open fields and grazing pastures with disturbed soils and open sunlight are
often more susceptible. Protecting grazing animals from toxic weeds in their primary feeding areas
is therefore important.

Control
Some guidelines to prevent the spread of noxious weeds are:

1. Avoid driving through noxious weed-infested areas.

2. Avoid transporting or planting seeds and plants that one can’t identify.

3. For noxious weeds in flower or with seeds on plants, pulling ‘gently’ out and placing in a
secure closable bag is recommended. Disposal such as hot composting or contained burn-
ing is done when safe and practical for the specific plant. Burning poison ivy can be fatal
to humans.

4. Using only certified weed-free seeds for crops or gardens.

Maintaining control of noxious weeds is important for the health of habitats, livestock, wildlife and
native plants, and of humans of all ages. How to control noxious weeds depends on the surround-
ing environment and habitats, the weed species, the availability of equipment, labor, supplies,
and financial resources. Laws often require that noxious weed control funding from governmental
agencies must be used for eradication, invasion prevention, or native habitat and plant community
restoration project scopes.

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10 Fundamentals of Weed Science

Noxious Weeds by Country


Australia
In Australia, the term “noxious weed” is used by state and territorial governments.

Canada
In Canada, constitutional responsibility for the regulation of agriculture and the environment are
shared between the federal and provincial governments. The federal government through the Ca-
nadian Food Inspection Agency (CFIA) regulates invasive plants under the authority of the Plant
Protection Act, the Seeds Act and statutory regulations. Certain plant species have been designat-
ed by the CFIA as noxious weeds in the Weed Seeds Order.

Each province also produces its own list of prohibited weeds. In Alberta, for example, a new Weed
Control Act was proclaimed in 2010 with two weed designations: “prohibited noxious” (46 spe-
cies) which are banned across Alberta, and “noxious” (29 species) which can be restricted at the
discretion of local authorities.

New Zealand
New Zealand has had a series of Acts of Parliament relating to noxious weeds: Noxious Weeds Act,
1908, Noxious Weeds Act 1950, and the Noxious Plants Act 1978. The 1978 Act was repealed by the
Biosecurity Act 1993 and words such as pest, organism and species are used in the new Act, rather
than “noxious”. Consequently, the term noxious weed is no longer used for official publications in
New Zealand.

United Kingdom
The Weeds Act, 1959 is described as “Preventing the spread of harmful or injurious weeds”, and
is mainly relevant to farmers and other rural settings rather than the allotment or garden-scale
grower. Five ‘injurious’ (that is, likely to be harmful to agricultural production) weeds are covered
by the provisions of the Weeds Act. These are:

• Spear thistle (Cirsium vulgare)

• Creeping, or field, thistle (Cirsium arvense)

• Curled Dock (Rumex crispus)

• Broad leaved dock (Rumex obtusifolius)

• Common ragwort (Senecio jacobaea)

The Department for Environment, Food and Rural Affairs (DEFRA) provides guidance for the
removal of these weeds from infested land. Much of this is oriented towards the use of herbicides.

The act does not place any automatic legal responsibility on landowners to control the weeds, but
they may be ordered to control them. Most common farmland weeds are not “injurious” within

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Types of Weed 11

the meaning of the Weeds Act and many such plant species have conservation and environmental
value. DEFRA has a duty to try to achieve reasonable balance among different interests. These
include agriculture, countryside conservation and the general public.

Section 14 of the Wildlife and Countryside Act 1981, makes it an offence to plant or grow certain
specified plants in the wild, including Giant Hogweed and Japanese Knotweed. Some local author-
ities have bye-laws controlling these plants. There is no statutory requirement for landowners to
remove these plants from their property.

United States
The federal government defines noxious weeds under the Federal Noxious Weed Act of 1974. Nox-
ious weeds are also defined by the state governments in the United States.

Abrus Precatorius
Abrus precatorius, known commonly as jequirity, Crab’s eye, or crab’s eye creeper, cock’s eyes,
rosary pea, paternoster pea, love pea, precatory pea or bean, prayer bead, John Crow Bead, coral
bead, red-bead vine, country licorice, Indian licorice, wild licorice, Jamaica wild licorice, Akar
Saga, coondrimany, gidee gidee, Jumbie bead ratti/rettee/retty, or weather plant, is a slender,
perennial climber that twines around trees, shrubs, and hedges. It is a legume with long, pin-
nate-leafleted leaves.

The plant is best known for its seeds, which are used as beads and in percussion instruments, and
which are toxic because of the presence of abrin. Ingestion of a single seed, well chewed, can be
fatal to both adults and children. The plant is native to India and grows in tropical and subtropical
areas of the world where it has been introduced. It has a tendency to become weedy and invasive
where it has been introduced.

Ecology and Invasiveness

Abrus precatorius from Koehler’s Medicinal-Plants

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12 Fundamentals of Weed Science

Abrus precatorius is a severely invasive plant in warm temperate to tropical regions, so much so that
it has become effectively pantropical in distribution. It had been widely introduced by humans, and
the brightly coloured and hard-shelled seeds had been spread by birds. By the end of the twentieth
century, it had been proclaimed as an invasive weed in many regions including some in Belize, Carib-
bean Islands, Hawaii, Polynesia and parts of the mainland United States. In Florida in particular, the
plant has invaded undisturbed pinelands and hammocks, including the vulnerable pine rocklands.

Once Abrus precatorius plants have grown to maturity under favourable conditions, their deep
roots are extremely difficult to remove, and the plants’ aggressive growth, hard-shelled seeds, and
ability to sucker, renders an infestation extremely difficult to eradicate and makes it very difficult
to prevent re-infestation. Herbicides such as glyphosate are effective, but need skilled application
if they are not to do more harm than good.

Toxin
The toxin abrin is a dimer consisting of two protein subunits, termed A and B. The B chain facil-
itates abrin’s entry into a cell by bonding to certain transport proteins on cell membranes, which
then transport the toxin into the cell. Once inside the cell, the A chain prevents protein synthesis
by inactivating the 26S subunit of the ribosome. One molecule of abrin will inactivate up to 1,500
ribosomes per second.

Symptoms are identical to those of ricin, except abrin is more toxic by almost two orders of mag-
nitude; the fatal dose of abrin is approximately 1/75th that of the fatal dose of ricin (though the
abrin in ingested seeds may be absorbed much more slowly than the ricin in Ricinus communis
even if the seeds are chewed and the coat penetrated, allowing time for successful rescue efforts in
at least some cases.) Abrin has an LD50 of only 0.56 μg/kg in mice, and Kingsbury lists a toxic dose
in humans at 0.00015% body weight, or approximately 0.1 mg for a 150 lb human. Ingesting intact
seeds may result in no clinical findings, as they can pass undigested through the gastrointestinal
tract because of their hard shell.

This plant is also poisonous to horses.

Symptoms of poisoning include nausea, vomiting, convulsions, liver failure, and death, usually
after several days.

Uses

The bright red seeds of A. precatorius are strung as jewellery.

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Types of Weed 13

Jewelry
The seeds of Abrus precatorius are much valued in native jewelry for their bright coloration. This
probably is unwise, mainly because young children are unfortunately drawn to the attractive beans
and may suck or chew on them. Most beans are black and red, suggesting a ladybug, though other
colors are available. Jewelry-making with jequirity seeds is somewhat hazardous. There are per-
sistent reports that the workers who pierce the seeds in order to thread them can suffer poisoning
or even death from a pinprick, but there seems to be little evidence. An online search found 265 sci-
entific papers referring to Abrus precatorius but not one of them dealt with occupational poisoning.

In Trinidad in the West Indies the brightly colored seeds are strung into bracelets and worn around
the wrist or ankle to ward off jumbies or evil spirits and “mal-yeux”—the evil eye. The Tamils use
Abrus seeds of different colors. The red variety with black eye is the most common, but there are
black, white and green varieties as well.

In March 2012 a recall was issued for bracelets made using Jequirity Beans sold by the Eden Proj-
ect and other outlets in the UK.

Unit of Measure
The seeds of Abrus precatorius are very consistent in weight. Formerly Indians used these seeds
to weigh gold using a measure called a Ratti, where 8 Ratti = 1 Masha; 12 Masha = 1 Tola (11.6
Grams).

As a Weapon
According to the 1898 King’s American Dispensatory,

Abrus seeds are the agents by which the Chamàr or “Native Skinner” caste of India carry on the
felonious poisoning of cattle for the purpose of securing their hides. This is done by means of small
spikes, called sui (needles) or sutari (awls), which are prepared by soaking the awl in a thin paste
of the water-soaked, pounded seeds, and then drying the weapon in the sun, after which it is oiled
and sharpened upon stone, affixed in a handle, and then used to puncture the skin of the animal.

An 1881 work by the District Superintendent of Police for British-occupied Bengal details the prepa-
ration and use of the sutari for the killing of cattle and in at least six murder cases. A native, prom-
ised a reduced sentence for the poisoning of a fellow villager’s bullock in exchange for his testimo-
ny, demonstrated the technique. First the outer shells of red or white seeds were cracked between
stones, then the two cotyledons from within thirty or forty seeds were soaked in water for ten min-
utes. These seeds were ground to a paste and rolled up into six sharp-ended one-inch cones, which
were inserted into either end of three pieces of straw and “exposed to the moderate influence of the
sun” to dry, whereupon they regained some of the original hardness of the seed. In this account, the
sutaries were described as being these cones, entirely made up of the hardened seed paste. The dried
cones were checked for sharpness, and if need be, whetted with a brick and re-set. Finally, to prevent
softening, they were waterproofed by “burying them for a night in some sort of animal grease.” For
testing, which Major Ramsay asked to be done exactly like a surreptitious killing, the prisoner set
two sutaries into a 1.5-inch wooden handle meant to be held in the hand by pressing them into a rag

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14 Fundamentals of Weed Science

stretched over sockets in the wood. A wandering “Brahmanee bull” was procured, and the prisoner
brought the sutari down in one direction and away in the other, so as to break off the cones inside the
animal’s flesh behind the horn, then pressed the skin over the broken ends leaving no obvious trace
of the injury. This process was repeated with two more cones to the base of the animal’s tongue. The
bull died after 34.5 hours, leaving no visible trace of the sutaries but a small amount of pus at the
wound site, whose swelling had mostly subsided by the time of death.

The 1890 Pharmacographia Indica gives an account, based in part on the above work, describing
the sutaris or suis (the terms being equivalent, depending on district, with the former based on the
object’s resemblance to the point of a cobbler’s awl). It describes the sutaris as 3/4 inch long and
weighing 1.5 to 2 grains, varying in color from dirty white to black, and describes the handle as 3
to 3.5 inches long and frequently made from two joints of bamboo wood, with sockets 1/4 to 3/8
inch deep and with the cavity exposed at one end for storage of additional sutaris. The weapons
were sometimes made with the milky juice of Calotropis gigantea instead of water, which was said
to speed the effect, and were sometimes supplemented with metallic mercury, dhatura, aconite,
and/or arsenic. It is added that “any attempt to withdraw the sutari by pulling at the pieces sticking
out, invariably breaks it, a portion being left in the wound.” One man murdered by a single blow
with a pair of sutaris died after three days; another, from whom the material had been successfully
excised, died three days later of tetanus. The price of one of these killings was said to total 16.5
rupees; the killers were punished by transportation for life.

In Traditional Medicine
Abrus precatorius, called kundu mani in Tamil and Guruvinda ginja in Telugu, has been used in
Siddha medicine for centuries. The white variety is used to prepare oil that is claimed to be an aph-
rodisiac. A tea is made from the leaves and used for fevers, coughs and colds. Seeds are poisonous
and therefore are used after mitigation. The Tamil Siddhars knew about the toxic effects in plants
and suggested various methods which is called “suththi seythal” or purification. This is done by
boiling the seeds in milk and then drying them. The protein is denatured when subjected to high
temperatures which removes its toxicity.

The plant is also used in Ayurveda and is said to promote hair growth. It is sometimes used as an
ingredient in Indian hair products.

Laboratory Study of Extracts


A variety of pharmacological effects have been observed in rodents, but have not been demonstrat-
ed clinically in humans, including:

• An ethanolic extract of Abrus precatorius was found to have antioxidant, anti-inflammato-


ry and analgesic potential in rodents.

• A methanolic extract of A. precatorius seeds causes reversible alterations in the estrous


cycle pattern and completely blocked ovulation in Sprague-Dawley rats.

• The methanolic extract produces dose-dependent bronchodilator activity in a guinea pig


model.

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Types of Weed 15

• Studies on aqueous, methanolic and chloroform extracts of Abrus precatorius showed


greater inhibitory activity against a number of disease-causing bacteria such as Bacillus
subtilis, Pseudomonas aeruginosa, Staphylococcus aureus, Salmonella typhimurium,
and Escherichia coli.

Names
Abrus precatorius has different names in various Indian and other languages.

• Arabic: ‫تيرفعلا نيع‬‎‎

• Badaga: gulugunji

• Assamese: Latumoni :(লাটুমণি)

• Bengali: Kunch: Koonch

• Gujarati: Gumchi: Chanothi

• Hebrew: Avrus ‎(‫)סורבא‬

• Hindi : Ratti, Gaungchi, Gunchi, Gunja

• Kannada: Gulaganji

• Kashmiri: Shangir

• Kreyòl ayisyen : Grenn Legliz, Lyann Legliz

• Indonesian: Saga rambat

• Malayalam: Kunni; Kunnikkuru കുന്നിക്കുരു

• Marathi: Gunj

• Nepali: Ratigedi

• Odia: Kaincha

• Persian: Gunchi; Chashami-Khurosa

• Punjabi : Mulati,Ratak,Labrigunchi

• Rajasthani : Chirmi

• Sanskrit: Gunja

• Sinhalese: Olinda

• Tamil: Gundumani, Kundumani

• Tagalog: Saga, Saga-saga, Saganamin

• Telugu: Gurivinda

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16 Fundamentals of Weed Science

• Yoruba: Ewe Omisinmisin or Ewe Ewerejeje

• Urdu: ‫جنگ‬‎

• Luo: Ombulu

• Zulu: Umkhokha

• Shangaan: Nsimani

• Ndebele: Amabope

Cultural Significance
In Rajasthan, India, Chirmi song is associated with this plant. There is also evidence that this plant has
significant economic value to the traditional Zulu people, due to the fact that it is a form of income for
unemployed Zulu people that make, and sell crafts that where made from the seeds of this plant.

Bromus Tectorum
Bromus tectorum, known as drooping brome or cheat grass, is a grass native to Europe, south-
western Asia and northern Africa, but has become invasive in many other areas.

Invasive Bromus tectorum on Spruce Mountain, Nevada

Description
Bromus tectorum is an annual bunchgrass, usually germinating in autumn, overwintering as a
seedling, then flowering in the spring or early summer. If winter rainfall is limiting and spring
moisture is adequate, the seeds will germinate in the spring, and the plants will flower that sum-
mer. It typically reaches 40–90 centimetres (16–35 in) tall, though plants as small as 2.5 centime-
tres (0.98 in) may produce seed. It is cleistogamous and self-fertile, with no evident out-crossing.

It has an extensive root system. The wide-spreading lateral roots are one of the keys to the survival
of this plant. A study showed that it had the capability to reduce soil moisture to the permanent
wilting point to a depth of 70 centimetres (28 in), reducing competition from other species.

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Types of Weed 17

Seeds
The seeds are dispersed by wind, small rodents, or attachment to animal fur, within a week of
maturity. They are also moved as a contaminant in hay, grain, straw, and machinery. It is an abun-
dant seed producer, with a potential in excess of 300 seeds per plant; seed production per plant is
dependent on plant density. Under optimal conditions, it may produce 450 kg of seed per hectare
(400 pounds per acre) with about 330,000 seeds/kg (150,000 seeds/pound).

The seeds maintain high viability in dry storage, lasting over 11 years. In the field, under buried
conditions, seeds will lose their viability in 2–5 years. The primary limit to germination is inade-
quate moisture. Seeds can withstand high soil temperatures. Germination is best in the dark or in
diffuse light. They germinate most quickly when covered with soil, but do not need to be in con-
tact with bare soil; some leaf litter cover will generally improve germination and establishment of
seedlings. Seedlings emerge rapidly from the top 2.5 cm (1”) of soil, and a few plants emerge from
depths of 8 cm (3”), but not from seeds 10 cm (4”) below the surface.

Habitat
Bromus tectorum grows in many climatic areas. It is found primarily in the 150–560 millimetres
(5.9–22.0 in) precipitation zone. It will grow in almost any type of soil, including B and C horizons
of eroded areas and areas low in nitrogen. It is most often found on coarse-textured soils and does
not grow well on heavy, dry, and/or saline soils. It grows in a relatively narrow range of soil tem-
peratures; growth starts at 2.0–3.5 °C (35.6–38.3 °F) and slows when temperatures exceed 15 °C
(59 °F).

Status as an Exotic Weed

Cheat grass in Elko, Nevada

Drooping brome has been introduced to southern Russia, west central Asia, North America, Ja-
pan, South Africa, Australia, New Zealand, Iceland, and Greenland. It was first found in the United
States (where it is known as downy brome or cheatgrass) in 1861 in New York and Pennsylvania,
by 1928 reaching throughout the United States (including Hawaii and Alaska), except for Florida
and portions of Alabama, Georgia, South Carolina; it is most abundant in the Great Basin, Colum-
bia Basin, and part of the introduced species that replaced California native plants in the Floristic
Province’s grasslands and other habitats.

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18 Fundamentals of Weed Science

Invasive Species
In the US, it grows on rangelands, pastures, prairies, fields, waste areas, eroded sites, and road-
sides. It is much reviled by ranchers and land managers. Drooping brome seeds are also a critical
portion of the diet of the chukar and grey partridge which have been introduced to the US. Inten-
sive sheep browsing of cheatgrass in early spring has been used as a fire fuels reduction strategy in
the hills adjacent to Carson City, Nevada.

The presence of biological soil crusts may slow the establishment of Bromus.

Drooping brome has demonstrated a quantitative and qualitative response to recent and near-
term changes in the concentration of atmospheric carbon dioxide. Laboratory experiments have
shown that above-ground biomass increased 1.5–2.7 gram per plant for every 10 part per million
(ppm) increase above the 270 ppm pre-industrial baseline. On the qualitative side, rising carbon
dioxide decreased the digestibility and potential decomposition of drooping brome. In addition
to stimulation of biomass, rising carbon dioxide may also increase the above ground retention
of drooping brome biomass by decreasing removal by animals or bacteria. Ongoing increases in
atmospheric carbon dioxide may contribute significantly to drooping brome productivity and fuel
load with subsequent effects on wildfire frequency and intensity.

B. tectorum has been shown to benefit from endophytic colonization by morels (Morchella sexte-
lata, M. snyderi) in western North America.

Cardoon
The cardoon (Cynara cardunculus), also called the artichoke thistle, cardone, cardoni, carduni, or
cardi, is a thistle-like plant in the sunflower family. It is a naturally occurring species that includes
the globe artichoke, and has many cultivated forms. It is native to the western and central Medi-
terranean region, where it was domesticated in ancient times.

Description
The wild cardoon is a stout herbaceous perennial plant growing 0.8 to 1.5  m (31 to 59  in) tall,
with deeply lobed and heavily spined green to grey-green tomentose (hairy or downy) leaves up to
50 cm (20 in) long, with yellow spines up to 3.5 cm long. The flowers are violet-purple, produced
in a large, globose, massively spined capitulum up to 6 cm (2 in) in diameter.

It is adapted to dry climates, native across an area from Morocco and Portugal east to Libya and
Greece and north to France and Croatia; it may also be native on Cyprus, the Canary Islands
and Madeira. In France, it only occurs wild in the Mediterranean south (Gard, Hérault, Aude,
Pyrénées-Orientales, Corsica). It has become an invasive weed in the pampas of Argentina, and is
also considered a weed in Australia and California.

In cultivation in the United Kingdom, this plant has gained the Royal Horticultural Society’s Award
of Garden Merit.

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Types of Weed 19

Cultivation
The two main cultivar groups are the cardoon (Cynara cardunculus Cardoon Group, syn. C. car-
dunculus var. altilis DC), selected for edible leaf stems, and the artichoke (Cynara cardunculus
Scolymus Group, sometimes distinguished as Cynara scolymus or C. cardunculus var. scolymus
(L.) Fiori), selected for larger edible flower buds. They differ from the wild plant in being larger
(up to 2 m tall), much less spiny, and with thicker leaf stems and larger flowers, all characteristics
selected by humans for greater crop yield and easier harvest and processing. Wild and cultivated
cardoons and artichokes are very similar genetically, and are fully interfertile, but only have very
limited ability to form hybrids with other species in the genus Cynara.

The earliest description of the cardoon may come from the fourth-century BC Greek writer Theo-
phrastus, under the name (Latin: cactus), although the exact identity of this plant is uncertain. The
cardoon was popular in Greek, Roman, and Persian cuisine, and remained popular in medieval
and early modern Europe. It also became common in the vegetable gardens of colonial America,
but fell from fashion in the late 19th century and is now very uncommon.

Cultivated cardoon foliage, Madrid Royal Botanical Garden, without leaf spines.

In Europe, cardoon is still cultivated in France (Provence, Savoie, Lyonnais), Spain, and Italy. In
the Geneva region, where Huguenot refugees introduced it about 1685, the local cultivar Argenté
de Genève (“Cardy”) is considered a culinary specialty. “Before cardoons are sent to table, the
stalks or ribs are blanched tying them together and wrapping them round with straw, which is
also tied up with cord, and left so for about three weeks”. Cardoons also are common vegetables in
northern Africa, often used in Algerian or Tunisian couscous.

Cardoon stalks can be covered with small, nearly invisible spines that can cause substantial pain if
they become lodged in the skin. Several spineless cultivars have been developed to overcome this.

Cardoon requires a long, cool growing season (about five months), but it is frost-sensitive. It also
typically requires substantial growing space per plant, so is not much grown except where it is
regionally popular.

Gastronomy
While the flower buds can be eaten much as small (and spiny) artichokes, more often the stems

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20 Fundamentals of Weed Science

are eaten after being braised in cooking liquid. Cardoon stems are for instance part of Lyonnaise
cuisine (cardoon au gratin). Only the innermost, white stalks are considered edible, and cardoons
are therefore usually prepared for sale by protecting the leaf stalks from the sunlight for several
weeks. This was traditionally done by burying the plant underground, thus, cardoon plantations in
Spain are often formed by characteristic earth mounds surrounding each plant, the earth covering
the stalks. In modern cultivation, the plant is usually instead wrapped in black plastic film or other
opaque material.

The flower buds of wild cardoons are still widely collected and used in southern Italy and Sicily. In
Spain and Portugal, the flower buds are also employed in cheesemaking: the pistils of the cardoon
flower are used as a vegetable rennet in the making of some cheeses such as the Torta del Casar and
the Torta de la Serena cheeses in Spain, or the azeitão cheese in Portugal.

Cardoon leaf stalks, which look like giant celery stalks, can be served steamed or braised, and have
an artichoke-like flavour with a hint of bitterness. They are harvested in winter and spring, being
best just before the plant flowers. In the Abruzzo region of Italy, Christmas lunch is traditionally
started with a soup of cardoon cooked in chicken broth with little meatballs (lamb or, more rarely,
beef), sometimes with the further addition of egg (which scrambles in the hot soup – called strac-
ciatella) or fried chopped liver and heart.

The cardoon stalks are considered a delicacy in Spain, particularly in the northern region of
Navarre, where they are grown in large quantities. In Spain, cardoons are typically cooked by
first boiling the stalks to soften them, and then adding simple sauces such as almond sauce
or small amounts of jamón; they are sometimes combined with clams, artichokes, or beans as
well.

Because of their seasonality (from November to February), cardoons are a staple of the Christmas
dinner in Navarre and the surrounding regions; for the same reason, cardoons are often sold as
vegetable preserves, usually in water or brine, so that they can be eaten all year round. Cardoons
are an ingredient in one of the national dishes of Spain, the cocido madrileño, a slow-cooking, one-
pot, meat and vegetable dish simmered in broth.

In the US, it is rarely found in conventional grocery stores but is available in some farmers’
markets in the months of May, June, and July. The main root can also be boiled and served
cold. The stems are also traditionally served battered and fried at St. Joseph’s altars in New
Orleans.

Cardoon is one of the herbs used to flavour Amaro liqueur, which may be called Cardamaro.

Other Uses
Cardoons are used as a vegetarian source of enzymes for cheese production. In Portugal, tradi-
tional coagulation of the curd relies entirely on this vegetable rennet. This results in cheeses such
as the Nisa (Denominazione di Origine Protetta), with a peculiar earthy, herbaceous and slightly
citric flavour that bears affinity with full-bodied or fortified wines.

The cardoon is also grown as an ornamental plant for its imposing architectural appearance, with
very bright silvery-grey foliage and large flowers in selected cultivars.

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Types of Weed 21

Cardoon has attracted recent attention as a possible source of biodiesel fuel. The oil, extracted
from the seeds of the cardoon, and called artichoke oil, is similar to safflower and sunflower oil in
composition and use.

Cardoon is the feedstock for the first biorefinery in the world converting the installations of a pet-
rochemical plant in Porto Torres, Sardinia, providing biomass and oils for the building blocks of
bioplastics. Matrica, the joint venture of ENI and Novamont, will inaugurate the facility in the Fall
of 2013.

Digitaria Ciliaris
Digitaria ciliaris is a species of grass known by the common names southern crabgrass, tropical
finger-grass, tropical crabgrass or summer grass.

The grass is known as “ගුරු තණ - guru thana” in Sri Lanka.

At Peradeniya Royal Botanical Garden.

Distribution
Digitaria ciliaris is a tough plant, believed to have originated in Asia but now found all over the
tropical belt of the planet, as well as in many temperate regions of both hemispheres. This grass is
an invasive species considered an aggressive weed in certain countries, including China, Mexico
and the United States.

Together with Portulaca oleracea, Ipomoea pes-caprae and Melanthera biflora, Digitaria ciliaris
is usually one of the first species colonizing degraded or altered environments in tropical zones of
the planet.

Description
This grass is as annual that can grow up to 1 m tall, but is usually much shorter. The roots are at
the nodes and the stems produce runners that allow the plant to grow fast forming scruffy-looking
patches about 1 m across and half a metre in height. The leaves are linear to linear-ovate narrowing
at the tip to 15 centimeters long. The inflorescence is at the top of a long stem, usually much taller
than the leaves, with two to nine 5–10 cm long sub-digitate racemes.

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22 Fundamentals of Weed Science

General appearance of the grass.


Plant and roots.

Juncus Effusus
Juncus effusus, with the common names common rush or soft rush, is a perennial herbaceous
flowering plant species in the family Juncaceae. In North America the common name soft rush
also refers to Juncus interior.

Distribution
Juncus effusus is nearly cosmopolitan, considered native in Europe, Asia, Africa, North America,
and South America. It has naturalized in Australia, Madagascar, and various oceanic islands.

It is found growing in wet areas, such as wetlands, riparian areas, and marshes. In the United
Kingdom it is found in purple moor-grass and rush pastures and fen-meadow plant associations.

Description

Pupal cases of Coleophora caespitiella on J. effusus.

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Types of Weed 23

Juncus effusus grows in large clumps about 1.5 metres (4 ft 11 in) tall at the water’s edge along
streams and ditches, but can be invasive anywhere with moist soil. It is commonly found growing
in humus-rich areas like marshes, ditches, fens, and beaver dams.

The stems are smooth cylinders with light pith filling. The yellowish inflorescence appears to
emerge from one side of the stem about 20 centimetres (8 in) from the top. In fact the stem ends
there; the top part is the bract, that continues with only a slight colour-band marking it from the
stem. The lower leaves are reduced to a brown sheath at the bottom of the stem.

Subspecies
Five subspecies are currently recognized:

1. Juncus effusus subsp. austrocalifornicus Lint — endemic to California and Baja California.

2. Juncus effusus subsp. effusus — widespread

3. Juncus effusus subsp. laxus (Robyns & Tournay) Snogerup — tropical Africa, Madagascar,
Mauritius, Canary Islands, Madeira.

4. Juncus effusus subsp. pacificus (Fernald & Wiegand) Piper & Beattie — Alaska, British
Columbia, Washington, Idaho, Oregon, California, Baja California.

5. Juncus effusus subsp. solutus (Fernald & Wiegand) Hämet-Ahti — central and eastern
United States.

Juncus effusus can be differentiated from the rarer Juncus pylaei by the number of ridges on the
stem. Juncus effusus has 30 to 40 ridges and J. pylaei has 10 to 20.

Chemistry

Juncusol is a 9,10-dihydrophrenathrene found in J. effusus. The plant also contains effusol and
dehydroeffusol.

Uses
Wildlife
The species provides wildfowl and wader feeding and nesting habitats, and also habitats for small
mammals.

A number of invertebrates feed on soft rush, including the rufous minor moth.

Humans
In Japan, this rush is grown to be woven into the covering of tatami mats. In Hui sup tea, Juncus
effusus is listed as one of the seven ingredients.

In Iran and Afghanistan too it is used to weave light cheap mat. It is called halfa (‫ )افلح‬and has
medicinal uses too.

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24 Fundamentals of Weed Science

In Europe, this rush was once used to make rushlights (by soaking the pith in grease), a cheap
alternative to candles.

Cultivation
The species is cultivated as an ornamental plant, for planting in water gardens, native plant and
wildlife gardens, and for larger designed natural landscaping and habitat restoration projects.

The cultivar Juncus effusus ‘Spiralis’ (syn. Juncus spiralis), with the common names corkscrew
rush or spiral rush, is a distinctive potted and water garden plant due to its very curled spiral like
foliage.

Weed Control
Juncus effusus can become a naturalized or invasive species, undesirable in rangelands for its
unpalatability to livestock. Suggested methods of controlling rushes include: ploughing; high ap-
plications of inorganic fertiliser (can pollute watersheds); and topping to prevent seed formation.

Hedera Helix
Hedera helix (common ivy, English ivy, European ivy, or just ivy) is a species of flowering plant in
the family Araliaceae, native to most of Europe and western Asia. A rampant, clinging evergreen
vine, it is a familiar sight in gardens, waste spaces, on house walls, tree trunks and in wild areas
across its native habitat. It is labeled as an invasive species in a number of areas where it has been
introduced.

Etymology
Hedera is the generic term for ivy. The specific epithet helix derives from Ancient Greek “twist,
turn”.

Synonyms
Synonyms include Hedera acuta, Hedera arborea (“tree ivy”), Hedera baccifera, Hedera grandi-
folia, bindwood, and lovestone.

Description
Hedera helix is an evergreen climbing plant, growing to 20–30  m (66–98  ft) high where suit-
able surfaces (trees, cliffs, walls) are available, and also growing as groundcover where no vertical
surfaces occur. It climbs by means of aerial rootlets with matted pads which cling strongly to the
substrate. The ability to climb on surfaces varies with the plants variety and other factors: Hedera
helix prefers non-reflective, darker and rough surfaces with near-neutral pH. It generally thrives
in a wide range of soil-pH with 6.5 being ideal, prefers moist, shady locations and avoids exposure
to direct sunlight, the latter promoting drying out in winter.

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Types of Weed 25

Ivy growing on a granite crag, Czech Republic.

Stems showing the rootlets used to cling to walls and tree trunks.

The leaves are alternate, 50–100 mm (2–4 in) long, with a 15–20 mm (0.6–0.8 in) petiole; they
are of two types, with palmately five-lobed juvenile leaves on creeping and climbing stems, and un-
lobed cordate adult leaves on fertile flowering stems exposed to full sun, usually high in the crowns
of trees or the top of rock faces.

The flowers are produced from late summer until late autumn, individually small, in 3-to-5 cm-di-
ameter (1.2-to-2.0 in) umbels, greenish-yellow, and very rich in nectar, an important late autumn
food source for bees and other insects.

The fruit are purple-black to orange-yellow berries 6–8 mm (0.2–0.3 in) in diameter, ripen-
ing in late winter, and are an important food for many birds, though somewhat poisonous to
humans.

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26 Fundamentals of Weed Science

One to five seeds are in each berry, which are dispersed after being eaten by birds.

The three subspecies are:

H. h. helix

central, northern and western Europe, plants without rhizomes, purple-black ripe fruit

H. h. poetarum Nyman (syn. Hedera chrysocarpa Walsh)

southeast Europe and southwest Asia (Italy, Balkans, Turkey), plants without rhizomes,
orange-yellow ripe fruit

H. h. rhizomatifera McAllister

southeast Spain, plants rhizomatous, purple-black ripe fruit

The closely related species Hedera canariensis and Hedera hibernica are also often treated as
subspecies of H. helix, though they differ in chromosome number so do not hybridise readily. H.
helix can be best distinguished by the shape and colour of its leaf trichomes, usually smaller and
slightly more deeply lobed leaves and somewhat less vigorous growth, though identification is
often not easy.

Range

Common ivy clinging on a London plane tree in Agde.

It ranges from Ireland northeast to southern Scandinavia, south to Portugal, and east to Ukraine
and Iran and northern Turkey.

The northern and eastern limits are at about the −2 °C (28 °F) winter isotherm, while to the west
and southwest, it is replaced by other species of ivy. Hedera helix itself is much more winter-hardy
and survives temperatures of −23.3 °C (−9.9 °F) (USDA Zone 6a) and above.

Cultivation and Uses


The ivy is widely cultivated as an ornamental plant. Within its native range, the species is greatly
valued for attracting wildlife. The flowers are visited by over 70 species of nectar-feeding insects,
and the berries eaten by at least 16 species of birds. The foliage provides dense evergreen shelter,
and is also browsed by deer.

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Types of Weed 27

Ivy-covered entrance to Malbork Castle.

In Europe, it is frequently planted to cover walls and the government recommends growing it on
buildings for its ability to cool the interior in summer, while providing insulation in winter, as well
as protecting the covered building from soil moisture, temperature fluctuations and direct expo-
sure to heavy weather. Further uses include weed suppression in plantings, beautifying unsightly
facades and providing additional green by growing on tree trunks.

Over 30 cultivars have been selected for such traits as yellow, white, variegated (e.g. ‘Glacier’),
and/or deeply lobed leaves (e.g. ‘Sagittifolia’), purple stems, and slow, dwarfed growth.

The following cultivars have gained the Royal Horticultural Society’s Award of Garden Merit:

• ‘Angularis aurea’       • ‘Glacier’ • ‘Midas Touch’

• ‘Caecilia’ • ‘Goldchild’ • ‘Parsley Crested’

• ‘Congesta’ • ‘Manda’s Crested’       • ‘Spetchley’

• ‘Duckfoot’

Ethnomedical Uses
Ivy extracts are part of current cough medicines. In the past, the leaves and berries were taken
orally as an expectorant to treat cough and bronchitis. In 1597, the British herbalist John Gerard
recommended water infused with ivy leaves as a wash for sore or watering eyes. The leaves can
cause severe contact dermatitis in some people. People who have this allergy (strictly a type IV hy-
persensitivity) are also likely to react to carrots and other members of the Apiaceae as they contain
the same allergen, falcarinol.

Invasive Species
Like other exotic species, ivy has predominantly been spread to areas by human action. H. helix is
labeled as an invasive species in many parts of the United States, and its sale or import is banned
in the state of Oregon.

Laurus nobilis and Ilex aquifolium are widespread relicts of the laurisilva forests that originally
covered much of the Mediterranean Basin when the climate of the region was more humid during
the tertiary era. Having disappeared during the glaciation, Ivy is believed to have been spread back

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28 Fundamentals of Weed Science

across the continent by birds once the continent warmed up again. With a great capacity for adap-
tation, Ivy will grow wherever development conditions and habitat similar to that of its European
origins exist, occurring as opportunistic species across a wide distribution with close vicariant
relatives and few species, indicating recent speciation.

Australia
It is considered a noxious weed across southern, particularly south-eastern, Australia and local
councils provide free information and limited services for removal. In some councils it is illegal to
sell the plant. It is a weed in the Australian state of Victoria.

Hedera in Hyde Park, Sydney used decoratively as underplanting

New Zealand
H. helix has been listed as an “environmental weed” by the Department of Conservation since
1990.

United States
In the United States, H. helix is considered weedy or invasive in a number of regions and is on the
official noxious weed lists in Oregon and Washington. Like other invasive vines such as kudzu,
H. helix can grow to choke out other plants and create “ivy deserts”. State- and county-sponsored
efforts are encouraging the destruction of ivy in forests of the Pacific Northwest and the Southern
United States. Its sale or import is banned in Oregon. Ivy can easily escape from cultivated gardens
and invade nearby parks, forests and other natural areas.

Control and Eradication


Ivy should not be planted or encouraged in areas where it is invasive. Where it is established, it is
very difficult to control or eradicate. In the absence of active and ongoing measures to control its
growth, it tends to crowd out all other plants, including shrubs and trees.

Damage to Trees
Ivy can climb into the canopy of young or small trees in such density that the trees fall over from

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Types of Weed 29

the weight, a problem which does not normally occur in its native range. In its mature form, dense
ivy can destroy habitat for native wildlife and creates large sections of solid ivy where no other
plants can develop.

Use as Building Facade Green


As with any self-climbing facade green, some care is required to make best use of the positive
effects: Ivy covering the walls of an old building is a familiar and often attractive sight. It has in-
sulating as well as weather protection benefits, dries the soil and prevents wet walls, but can be
problematic if not managed correctly.

Ivy, and especially European ivy (H. helix) grows vigorously and clings by means of fibrous roots,
which develop along the entire length of the stems. These are difficult to remove, leaving an un-
sightly “footprint” on walls, and possibly resulting in expensive resurfacing work. Additionally, ivy
can quickly invade gutters and roofspaces, lifting tiles and causing blockages. It also harbors mice
and other unwelcome creatures. The plants have to be cut off at the base, and the stumps dug out
or killed to prevent regrowth.

Therefore, if a green facade is desired, this decision has to be made consciously, since later removal
would be tedious.

Kudzu

Flowers of Pueraria montana

Kudzu is a group of plants in the genus Pueraria, in the pea family Fabaceae, subfamily Faboideae.
They are climbing, coiling, and trailing perennial vines native to much of eastern Asia, Southeast
Asia, and some Pacific islands. The name comes from the Japanese name for the plants, kuzu (クズ
or 葛?), which was written “kudzu” in historical romanizations. Where these plants are naturalized,
they can be invasive and are considered noxious weeds. The plant climbs over trees or shrubs and

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30 Fundamentals of Weed Science

grows so rapidly that it kills them by heavy shading. The plant is edible, but often sprayed with
herbicides.

Taxonomy and Nomenclature

Kudzu seedpods

The name kudzu describes one or more species in the genus Pueraria that are closely related, and
some of them are considered to be varieties rather than full species. The morphological differences
between them are subtle; they can breed with each other, and introduced kudzu populations in the
United States apparently have ancestry from more than one of the species. They are:
• P. montana
• P. lobata (P. montana var. lobata)
• P. edulis
• P. phaseoloides
• P. thomsonii (P. montana var. chinensis)
• P. tuberosa

Origin in The United States


Kudzu was introduced to the United States as an ornamental bush and an effortless and efficient
shade producer at the Philadelphia Continental Exposition in 1876. In the 1930s and ‘40s, the vine
was rebranded as a way for farmers to stop soil erosion. Workers were paid eight dollars an hour to
sow topsoil with the invasive vine. The cultivation covered over one million acres of kudzu.

Propagation
Kudzu spreads by vegetative reproduction via stolons (runners) that root at the nodes to form new
plants and by rhizomes. Kudzu will also spread by seeds, which are contained in pods and mature

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Types of Weed 31

in the autumn, although this is rare. One or two viable seeds are produced per cluster of pods. The
hard-coated seeds can remain viable for several years, and will successfully germinate only when
soil is persistently soggy for five to seven days, with temperatures above 20 °C (68 °F). Once ger-
minated, saplings must be kept in a well-drained medium that retains high moisture. During this
stage of growth, it is critical for kudzu to receive as much sunlight as possible. Kudzu saplings are
sensitive to mechanical disturbance, and are damaged by chemical fertilizers. They do not tolerate
long periods of shade, or high water tables.

Uses
Soil Improvement and Preservation
Kudzu has been used as a form of erosion control and also to enhance the soil. As a legume, it in-
creases the nitrogen in the soil by a symbiotic relationship with nitrogen-fixing bacteria. Its deep
taproots also transfer valuable minerals from the subsoil to the topsoil, thereby improving the top-
soil. In the deforested section of the central Amazon Basin in Brazil, it has been used for improving
the soil pore-space in clay latosols, thus freeing even more water for plants than in the soil prior
to deforestation.

Animal Feed
Kudzu can be used by grazing animals, as it is high in quality as a forage and palatable to live-
stock. It can be grazed until frost and even slightly after. Kudzu had been used in the southern
United States specifically to feed goats on land that had limited resources. Kudzu hay typically
has a 15–18% crude protein content and over 60% total digestible nutrient value. The quality
of the leaves decreases, however, as vine content increases relative to the leaf content. Kudzu
also has low forage yields despite its rate of growth, yielding around two to four tons of dry
matter per acre annually. It is also difficult to bale due to its vining growth and its slowness in
shedding water. This makes it necessary to place kudzu hay under sheltered protection after
being baled. Kudzu is readily consumed by all types of grazing animals, yet frequent grazing
over three to four years can ruin stands. Thus, kudzu only serves well as a grazing crop on a
temporary basis.

Basketry
Kudzu fiber has long been used for fiber art and basketry. The long runners which propagate
the kudzu fields and the larger vines which cover trees make excellent weaving material. Some
basketmakers use the material green. Others use it after splitting it in half, allowing it to dry
and then rehydrating it using hot water. Both traditional and contemporary basketry artists
use kudzu.

Medicine
Kudzu contains a number of useful isoflavones, including puerarin, about 60% of the total isofla-
vones, and also daidzein (an anti-inflammatory and antimicrobial agent) and daidzin (structurally
related to genistein). It has shown value in treating migraine and cluster headaches.It is recom-
mended for allergies and diarrhea.

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32 Fundamentals of Weed Science

Kudzu leaves near Canton, Georgia

In traditional Chinese medicine, where it is known as gé gēn (Chinese: 葛根), kudzu is considered
one of the 50 fundamental herbs. It is used to treat tinnitus, vertigo, and Wei syndrome (superfi-
cial heat).

Kudzu has been used as a remedy for alcoholism and hangover. The root was used to prevent ex-
cessive consumption, while the flower was supposed to detoxify the liver and alleviate the symp-
toms afterwards. However, a 2007 study suggested that the use of the kudzu root is inappropriate
as a hangover remedy due to increased acetaldehyde accumulation through mitochondrial alde-
hyde dehydrogenase inhibition. Some hangover remedies are marketed with kudzu as one of their
active ingredients.

It has also shown potential in animal models of Alzheimer’s disease.

Food and Beverages

Kuzumochi (葛餅), Japanese-style kudzu starch cake (Katori City, Japan)

The roots contain starch, which has traditionally been used as a food ingredient in East Asia. In
Vietnam, the starch called bột sắn dây is flavoured with pomelo oil and then used as a drink in the
summer. In Japan, the plant is known as kuzu and the starch named kuzuko. Kuzuko is used in
dishes including kuzumochi, mizu manjū, and kuzuyu. It also serves as a thickener for sauces, and
can substitute for cornstarch.

The flowers are used to make a jelly that tastes similar to grape jelly. Roots, flowers, and leaves of
kudzu show antioxidant activity that suggests food uses. Nearby bee colonies may forage on kudzu

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Types of Weed 33

nectar during droughts as a last resort, producing a low-viscosity red or purple honey that tastes
of grape jelly or bubblegum.

Kudzu has also been used for centuries in East Asia to make herbal teas and tinctures. Kudzu pow-
der is used in Japan to make an herbal tea called kuzuyu.

Other Uses
Kudzu fiber, known as ko-hemp, is used traditionally to make clothing and paper, and has also
been investigated for industrial-scale use. The stems are traditionally used for basketry.

It may become a valuable asset for the production of cellulosic ethanol. In the Southern United
States, kudzu is used to make soaps, lotions, and compost.

Invasive Species

Kudzu growing on trees in Georgia, United States

Kudzu plants near Canton, Georgia

Ecological Damage and Roles


Kudzu’s environmental and ecological damage results from acting through “interference competi-
tion”, meaning it outcompetes other species for a resource. Kudzu competes with native flora for
light, and acts to block their access to this vital resource by growing over them and shading them
with its leaves. Native plants may then die as a result.

Changes in leaf litter associated with kudzu infestation results in changes to decomposition pro-
cesses and a 28% reduction in stocks of soil carbon, with potential implications for processes in-
volved in climate change.

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34 Fundamentals of Weed Science

United States
Kudzu was introduced from Japan into the United States at the Japanese pavilion in the 1876 Cen-
tennial Exposition in Philadelphia. It is now common along roadsides and other disturbed areas
throughout most of the southeastern United States as far north as rural areas of Pulaski County,
Illinois. Estimates of its rate of spreading differ wildly; it has been described as spreading at the
rate of 150,000 acres (610 km2) annually, although in 2015 the United States Forest Service esti-
mated the rate to be only 2,500 acres per year.

Canada
Kudzu was discovered July 2009 in a patch 110  m (360  ft) wide and 30  m (98  ft) across, on a
south-facing slope on the shore of Lake Erie near Leamington, Ontario, about 50 km (31 mi) south-
east of Windsor. Leamington is located in the second-warmest growing region of Canada after
south coastal British Columbia.

Ecologist Gerald Waldron made the Leamington find while walking along the beach. He recognized
the kudzu instantly, having read about its destructive expansion in the southeastern United States.

Other Countries
During World War II, kudzu was introduced to Vanuatu and Fiji by United States Armed Forces to
serve as camouflage for equipment and has become a major weed.

Kudzu is also becoming a problem in northeastern Australia, and has been seen in isolated spots
in Northern Italy (Lake Maggiore).

In New Zealand, kudzu was declared an “unwanted organism” and was added to the Biosecurity
New Zealand register in 2002.

Control
Crown Removal
For successful long-term control of kudzu, it is not necessary to destroy the underground
system, which can be extremely large and deep. It is only necessary to use some method to
kill or remove the kudzu root crown and all rooting runners. The root crown is a fibrous knob
of tissue that sits on top of the roots. Crowns form from multiple vine nodes that root to the
ground, and range from pea- to basketball-sized. The older the crowns, the deeper they tend to
be found in the ground. Nodes and crowns are the source of all kudzu vines, and roots cannot
produce vines. If any portion of a root crown remains after attempted removal, the kudzu plant
may grow back.

Mechanical methods of control involve cutting off crowns from roots, usually just below ground
level. This immediately kills the plant. Cutting off the above-ground vines is not sufficient for an
immediate kill. It is necessary to destroy all removed crown material. Buried crowns can regen-
erate into healthy kudzu. Transporting crowns in soil removed from a kudzu infestation is one
common way that kudzu unexpectedly spreads and shows up in various locations.

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Types of Weed 35

Mowing
Close mowing every week, regular heavy grazing for many successive years, or repeated cultivation
may be effective, as this serves to deplete root reserves. If done in the spring, cutting off vines must
be repeated. Regrowth appears to exhaust the plant’s stored carbohydrate reserves. Cut kudzu can
be fed to livestock, burned, or composted.

Grazing
The city of Chattanooga, Tennessee, has undertaken a trial program using goats and llamas to
graze on the plant. Similar efforts to reduce widespread nuisance kudzu growth have also been
undertaken in the cities of Winston-Salem, North Carolina and Tallahassee, Florida.

Fire
Prescribed burning is also used on old extensive infestations to remove vegetative cover and promote
seed germination for removal or treatment. While fire is not an effective way to kill kudzu, equipment,
such as a skid loader, can later remove crowns and thereby kill kudzu with minimal disturbance of soil.

Herbicide
A systemic herbicide, for example, glyphosate, Triclopyr, or Tordon, can be applied directly on cut
stems, which is an effective means of transporting the herbicide into the kudzu’s extensive root
system. Herbicides can be used after other methods of control, such as mowing, grazing, or burn-
ing, which can allow for an easier application of the chemical to the weakened plants. In large-scale
forestry infestations, soil-active herbicides have been shown to be highly effective.

After initial herbicidal treatment, follow-up treatments and monitoring are usually necessary, de-
pending on how long the kudzu has been growing in the area. It may require up to 10 years of su-
pervision after the initial chemical placement to make sure the plant does not return.

Fungi
Since 1998, the United States Department of Agriculture, Agricultural Research Service (ARS)
has experimented with using the fungus Myrothecium verrucaria as a biologically based herbi-
cide against kudzu. A diacetylverrucarol spray based on M. verrucaria works under a variety of
conditions (including the absence of dew), causes minimal injury to many of the other woody
plants in kudzu-infested habitats, and takes effect quickly enough that kudzu treated with it in the
morning starts showing evidence of damage by midafternoon. Initial formulations of the herbicide
produced toxic levels of other trichothecenes as byproducts, though the ARS discovered growing
M. verrucaria in a fermenter on a liquid instead of a solid diet limited or eliminated the problem.

Beneficial Weed
A beneficial weed is a plant not generally considered domesticated and often viewed as a weed
but which has some companion plant effect, is edible, contributes to soil health, or is otherwise

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36 Fundamentals of Weed Science

beneficial. Beneficial weeds include many wildflowers, as well as other weeds that are commonly
removed or poisoned.

Clover was once included in grass seed mixes, because it is a legume that fertilizes the soil

Soil Health

Dandelions benefit neighboring plant health by bringing up nutrients and moisture with its deep tap root

Although erroneously assumed to compete with neighboring plants for food and moisture, some
“weeds” provide the soil with nutrients, either directly or indirectly.

• For example, legumes, such as white clover, if they are colonized by the right bacteria (Rhi-
zobium most often) add nitrogen to the soil through the process of nitrogen fixation, where
the bacteria has a symbiotic relationship with its hosts roots, “fixing” atmospheric nitrogen
(combining it with oxygen or hydrogen) making the nitrogen plant-available (NH4 or NO3).

• Others use deep tap roots to bring up nutrients and moisture from beyond the range of
normal plants so that the soil improves in quality over generations of that plant’s presence.

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Types of Weed 37

• Weeds with strong, widespread roots also introduce organic matter to the earth in the form
of those roots, turning hard, dense clay dirt into richer, more fertile soil.

• Some plants like tomatoes and corn will “piggyback” on nearby weeds, allowing their rela-
tively weak root systems to go deeper.

Pest Prevention
Many weeds protect nearby plants from insect pests.

Some beneficial weeds repel insects and other pests through their smell , for example alliums and
wormwood. Some weeds mask a companion plant’s scent, or the pheromones of pest insects, as
with ground ivy, as well as oregano and other mints.

Some also are unpleasant to small animals and ground insects, because of their spines or other
features, keeping them away from an area to be protected.

Crow garlic, like any allium, masks scents from pest insects, protecting neighboring plants

Trap Crops
Some weeds act as trap crops, distracting pests away from valued plants. Insects often search for
target plants by smell, and then land at random on anything green in the area of the scent. If they
land on an edible “weed”, they will stay there instead of going on to the intended victim. Some-
times, they actively prefer the trap crop.

Host-finding Disruption
Recent studies on host-plant finding have shown that flying pests are far less successful if their
host-plants are surrounded by any other plant or even “decoy-plants” made of green plastic, card-
board, or any other green material.

• First, they seek plants by scent. Any “weed” that has a scent reduces the odds of them find-
ing crop plants. Examples are Crow Garlic (wild chives) and Ground Ivy (a form of wild
mint), both dramatically masking both plant scent and insect pheromones. They cut down

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38 Fundamentals of Weed Science

Japanese beetle infestation, and caterpillar infestation, for example cabbage worm, tomato
hornworm, and even squash bugs.

• Second, once an insect is near its target, it avoids landing on dirt, but lands on the nearest
green thing. Bare earth gardening helps them home in perfectly on the victim crop. But if
one is using “green mulch”, even grass or clover, the odds are that they will make what’s
called an “inappropriate landing” on some green thing they don’t want. They will then fly a
short distance at random, and land on any other green thing. If they fail to accidentally hit
the right kind of plant after several tries, they give up.

• If they plan to lay eggs on the crop, weeds provide one more line of defense: Even if they
find the right plant, in order to ensure that they didn’t hit on a dying plant or falling
leaf, they then make short leaf-to-leaf flights before laying eggs. They must land on the
“right kind of leaf” enough times in sequence, before they will risk laying their eggs.
The more other greenery is nearby, the harder it is for them to remain on target and
get enough reinforcement. Enough “inappropriate landings”, and they give up, heading
elsewhere.

One scientific study said that simply having clover growing nearby cut the odds of cabbage root
flies hitting the right plant from 36% to 7%.

Companion Plants

Queen Anne’s Lace provides shelter to nearby plants, as well as attracting predatory insects that eat pests like
caterpillars, and may boost the productivity of tomato plants

Many plants can grow intercropped in the same space, because they exist on different levels in the
same area, providing ground cover or working as a trellis for each other. This healthier style of
horticulture is called forest gardening. Larger plants provide a wind break or shelter from noonday
sun for more delicate plants.

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Types of Weed 39

Green Mulch
Conversely, some intercropped plants provide living mulch effect, used by inhibiting the growth
of any weeds that are actually harmful, and creating a humid, cooler microclimate around nearby
plants, stabilizing soil moisture more than they consume it for themselves.

Plants such as ryegrass, red clover, and white clover are examples of “weeds” that are living mulch-
es, often welcomed in horticulture.

Herbicide
Repel plants or fungi, through a chemical means known as allelopathy. Specific other plants can be
bothered by a chemical emission through their roots or air, slowing their growth, preventing seed
germination, or even killing them.

Beneficial Insects
A common companion plant benefit from many weeds is to attract and provide habitat for benefi-
cial insects or other organisms which benefit plants.

For example, wild umbellifers attract predatory wasps and flies. The adults eat nectar, but they
feed common garden pests to their offspring .

Some weeds attract ladybugs or the “good” types of nematode, or provide ground cover for pred-
atory beetles.

Uses for Humans


• Some beneficial weeds, such as lamb’s quarters and purslane, are edible. This list of edible
flowers includes many wildflowers that are considered weeds when not planted intention-
ally. Dandelion is an example of an edible weed.
• A number of weeds have been proposed as natural alternate sources for latex (rubber),
including goldenrod, from which the tires were made on the car famously given by Henry
Ford to Thomas Edison.
• Cocklebur and stinging nettle have been used for natural dyes.
• Milkweed is a more effective insulator than goose down.
• Some plants seem to subtly improve the flavor of other plants around them, for example
stinging nettle, besides being edible if properly cooked, seems to increase essential oil pro-
duction in nearby herbs.

Examples
• Clover is a legume. Like other beans, it hosts bacteria that fix nitrogen in the soil. Its vining
nature covers the ground, sheltering more moisture than it consumes, providing a humid,
cooler microclimate for surrounding plants as a “green mulch”. It also is preferred by ro-
dents over many garden crops, reducing the loss of vegetable crops.

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40 Fundamentals of Weed Science

• Dandelions possess a deep, strong tap root that breaks up hard soil, benefiting weaker-root-
ed plants nearby, and draw up nutrients from deeper than shallower-rooted nearby plants
can access. They will also excrete minerals and nitrogen through their roots.
• Crow garlic, the wild chives found in sunny parts of a North American yard, has all of the
companion plant benefits of other alliums, including repelling japanese beetles, aphids,
and rodents, and being believed to benefit the flavor of solanums like tomatoes and pep-
pers. It can be used as a substitute for garlic in cooking, though it may lend a bitter after-
taste.
• Bishop’s lace (Queen Anne’s Lace) works as a nurse plant for nearby crops like lettuce,
shading them from overly intense sunlight and keeping more humidity in the air. It attracts
predatory wasps and flies that eat vegetable pests. It has a scientifically tested beneficial
effect on nearby tomato plants. When it is young it has an edible root, revealing its relation-
ship to the domesticated carrot.

List of Beneficial Weeds


This is a list of undomesticated or feral plants, generally considered weeds, yet having some posi-
tive effects or uses, often being ideal as companion plants in gardens.

Beneficial weeds can accomplish a number of roles in the garden or yard, including fertilizing the
soil, increasing moisture, acting as shelter or living mulch, repelling pests, attracting beneficial
insects, or serving as food or other resources for human beings.

Chart

Beneficial Weed Chart

Common Scientific Companion Attracts/


Repels Traps Edibility Medicinal Avoid Comments
name name plant for hosts

Its extract
immobilizes the
filariform larvae
of Strongyloides
stercoralis in less
than one hour.
In contempo-
rary medicine,
Mimosa pudica is
Used as
ground being investigat-
a natural
Bashful Mimosa cover for predatory ed for its potential
ground
Mimosa pudica tomatoes, beetles to yield novel
cover in
peppers chemotherapeu-
agriculture
tic compounds.
It contains an
alkaloid called
mimosine, which
has been found
to have potent
antiproliferative
and apoptotic
effects.

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Types of Weed 41

Many
domesticat-
Used in folk
ed animals
Caper Euphorbia medicine as an
Moles can eat it,
Spurge lathyris antiseptic and
although it is
purgative
poisonous to
humans.

Brassica
(cabbage
and cousins
like broc-
This legume
coli and
hosts nitro-
cauliflow-
gen-fixing
er), corn,
bacteria in
cucurbits
This its roots, and
(cucumber,
legume is a Night- therefore
squash,
high-pro- shades fertilizes
melons,
tein source (to- the soil for
gourds) --
Clover Trifolium Rabbits of food, but mato, neighboring
Along with
generally pepper, plants. It is
fertilizing
only eaten egg- also used
the soil,
in survival plants) as a fallow
this plant
situations plant by
provides a
some farm-
humid mi-
ers, and is a
croclimate
very popular
that benefits
fodder plant.
many plants
by stabi-
lizing their
moisture

Poison-
ous to Also used
Grasses Army Is used in Chi-
Cocklebur Xanthium some for yellow
and grains worms nese medicine
life- dye
stock

Common Scientific Companion Attracts/


Repels Traps Edibility Medicinal Avoid Comments
name name plant for hosts

fruit trees,
nightshades 3-mercap-
(tomatoes, to-2-methyl-
peppers, slugs, pentan-1-ol
Can be This is a wild
potatoes, aphids, in onion was beans,
Crow Allium used like cousin of
etc.), carrot fly, found to have an peas,
garlic vineale convention- onions and
brassicas cabbage antioxidant potent parsley
al chives garlic
(cabbage, worms that inhibits per-
broccoli, oxynitrite induced
kohlrabi, diseases.
etc.) carrots

Tap root
breaks up
hardened
soil and
brings up
In season, nutrients
Various Used as a
Army- leaves and from deep
Dandelion Taraxacum grains, to- Honeybees diuretic in herbal
worms flowers are down, bene-
mato plants medicine
edible fiting plants
with weaker
or shallower
roots without
competing
with them.

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42 Fundamentals of Weed Science

Contains
latex, the
automobile
Pear trees, given to
Black Various Thomas
Predatory Numerous medic-
Goldenrod Solidago Locust Lepidop- Edison by
wasps inal uses
Tree, Sugar tera larvae Henry Ford
Maple had tires
made from
goldenrod
latex

This wild
mint makes
Used in the tra- a good
Tomatoes,
ditional medicine ground
peppers, cabbage
of Europe going cover com-
cucum- worms,
back thousands panion plant,
bers and cucumber
of years. Inflam- creating a
relatives worms Can be
Glechoma mation of the humid mi-
Ground Ivy (squash, and bee- used in
hederacea eyes, tinnitus, a croclimate,
melons), tles, to- herbal teas
diuretic, astrin- covering up
broccoli, mato horn
gent, tonic and nearby plant
brussels worms,
gentle stimulant. scents, and
sprouts, others
distracting
cauliflower
pests from
companion
crops.

Common Scientific Companion Attracts/


Repels Traps Edibility Medicinal Avoid Comments
name name plant for hosts

The berries Ripe fruit, when


of this fruit cooked, is used
Horsen- Solanum Predatory
may be ed- by herbalists as
ettle carolinense beetles
ible when a diuretic and
cooked sedative

Can be used
as a more
effective
insulator
than goose
down. Emits
a chemical
that breaks
Predatory Folk remedy up hard soil,
Corn, basil, wasps and Wire- for warts, sap allowing
Milkweed Asclepias
potatoes the Monarch worms reduces poison nearby
butterfly ivy symptoms plants to
develop
healthier
root sys-
tems. Basil
repels some
insects
that attack
milkweed.

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Types of Weed 43

Most
vegetables,
Considered
especially
one of the
brassica
Squash “magic
(cabbage, All parts of
bugs, bullet”
broccoli, this plant
cucumber trap crop companion
et al.), are edible,
beetles, for cat- plants, bene-
cucurbits Predatory flowers
Nasturtium Tropaeolum striped erpillars fiting almost
(cucumbers, wasps and leaves
pumpkin and black any crops
melons, make bril-
beetles, aphids around it in
squash) and liant salad
woolly some way,
solanum decoration
aphids and not
(tomatoes,
known to
potatoes,
hurt any
peppers,
etc.)

Also once
grown as
a crop for
its fiber. Its
Despite
juice was
its “sting”,
once used
young plant
in the place
parts are
of rennet in
edible, as is One of the most-
broccoli, cheese-mak-
much of the used plants in
Urtica tomato , Va- ing. It
Nettle plant when herbal medicine,
dioica lerian, mint, was also
blanched or with a long list of
fennel a source
otherwise benefits
of “green”
prepared.
for dye. It
Also makes
can still be
a nutritious
used as a
herbal tea
high-protein
additive in
animal feed,
once dried.

Common Scientific Companion Attracts/


Repels Traps Edibility Medicinal Avoid Comments
name name plant for hosts

Purslane
is eaten
throughout
much of
Europe and Breaks up
In Traditional
Mexico. It hard soil and
Chinese Medi-
contains hardpan,
cine, it is used to
more brings
treat infections or
Omega-3 nutrients and
bleeding of the
corn, sola- fatty acids Dill, water up
genito-urinary
Portulaca nums like than any pars- from deeper
Purslane tract as well as
oleracea tomatoes other leafy nip, than crops
dysentery. It may
and peppers vegetable radish can reach,
also be applied
plant. It can provides
topically to relieve
be eaten healthy
sores and insect
in salad, ground cov-
or snake bites on
stir-fried, or er, stabilizing
the skin.
cooked like soil moisture
spinach.
berries can
be eaten
like capers

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44 Fundamentals of Weed Science

Nightshades
(especially Some recent Do not con-
Dill,
Queen tomatoes), predatory Young scientific support fuse with its
Daucus pars-
Anne’s alliums wasps and roots are for its historic poisonous
carota nip,
Lace (onions, flies edible use as a herbal cousin, wa-
radish
chives), contraceptive ter hemlock
lettuce

Domesticat-
ed mustard
is a hybrid of
Grape vine three differ-
, radish, Traps ent species
non-mus- various of wild
Seeds and
Wild Mus- Brassica- tard brassi- brassica mustard, all
Ladybugs leaves are beets
tard ceae ca, including pests, of which are
edible
cabbage, including still used in
cauliflower, aphids some places
broccoli for food. This
is known as
the Triangle
of U.

This
includes the
feral multi-
flora rose,
brought to
the US both
for use as
root stock for
domesticat-
Rose hips
Strawber- Traps Same medicinal ed roses,
Rodents can be
Wild Rose Rosa ries, grapes, Japanese benefits as do- and as a
and deer used in
roses beetles mesticated rose “natural
herbal teas
fence” for
lifestock.
In the mid
20th century
miles of mul-
tiflora rose
hedge were
planted in
sequence.

Common Scientific Companion Attracts/


Repels Traps Edibility Medicinal Avoid Comments
name name plant for hosts

This legume
fixes nitro-
gen, allow
Pepper
it to grow
and tomato
in a tomato
plants,
garden only
brassica Provides
until time
(cabbage, ground
Vicia ameri- to plant, as
Wild Vetch mustard, cover for
cana ground cov-
broccoli), predatory
er. But can
other plants beetles
be left grow-
needing
ing among
high nitro-
brassica for
gen
additional
nitrogen and
microclimate

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Types of Weed 45

Categories of beneficial weeds


Pest-repellent
• Neem—repels leaf eating insects

Edible
• Borage—all parts have various medicinal purposes, with the edible flowers being used in
desserts.

• Burdock—roots are edible (as are the stalks, but particularly the young leaves )

• Chickweed (Stellaria media) -- used in salads and also as ground cover

• Cornflower various colours; can be served as edible garnish to decorate salads.

• Horsetail—primeval plant that produces its own vitamin D & is high in silica; tops are very
similar to & may be eaten like asparagus

• Lamb’s quarters—leaves and shoots, raw, also prevents erosion, also distracts leaf miners
from nearby crops

• Nettle—young leaves collected before flowering used as a tea or spinach substitute. Plants
have use as compost material or for fibre.

• Purslane—prepared raw for salads or sautéed

• Shepherd’s purse—leaves are edible and often sautéed or blanched

• Watercress—can be eaten raw or cooked; is considered a weed in some cultures (caution


required when harvesting wild due to the risk of contracting potentially fatal liver fluke)

Habitat for Beneficial Insects


• Wild blackberry—attracts predatory insects, and produces berries

• Motherwort—attracts bees

• Joe-Pye weed—habitat for pollinators and predatory insects

• Aster—habitat predatory insects

Shelter Plants
• Normal grass can be used as ground cover, especially in nitrogenous soils.

Trap Crops
Trap crops draw potential pests away from the actual crop intended for cultivation.

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46 Fundamentals of Weed Science

• Cowpea—attracts ladybird beetle, so planting around cotton fields protects them from
sucking insects. It serves as source of food and niche.

Medicinal Use
• Rumex—Dock. Commonly grows in association with nettle, is rumoured to cure or ease
their sting. Crush a leaf before applying to affected area. Liver tonic, increases iron levels in
the blood, regulates hormone levels and reduces menstrual flow and cramping.
• Urtica dioica—Stinging nettle. Nutritive tonic, diuretic. Restorative to the adrenals and
kidneys. Highly nutritive. Can be eaten after cooking to remove the stinging properties.
Stinging nettle has been used as a successful pain reliever for arthritis by stinging the af-
fected area with the raw plant.
• Arctium lappa—Burdock. The root is used to stimulate detoxification of the lymph and
liver, known as a “blood purifier.” It also has diuretic and diaphoretic properties.
• Taraxacum officinale—Dandelion. The leaves are a tonic to the kidneys, being one of the
few diuretics that does not deplete the body of potassium. The whole plant, especially the
root, is a detoxifying tonic for the liver. The whole plant is bitter and can be used as a di-
gestive stimulant.
• Capsella bursa-pastoris—Shepherd’s Purse. Used to stop bleeding.
• Galium aparine – used in folk medicine to make poultices.
• Stellaria media -- Chickweed. One of the most nutrient dense plants, full of antioxidants.
Effective for skin conditions when infused into oil. Dissolves cysts and lumps.
• Plantago spp—Plantain, Ribwort, Pig’s ear. Wound healing herb. “Not only does plan-
tain increase the speed of healing, it also relieves pain, stops bleeding, draws out for-
eign matter, stops itching, prevents and stops allergic reactions from bee stings, kills
bacteria, and reduces swelling.” Mucilaginous. The seed husks are the main ingredient
in psyllium laxatives. Identify this common weed by the 5 parallel veins on the under-
side of the leaf.
• Hypericum perforatum—St John’s Wort. Tincture is used as an anti-viral and for muscle
aches, shingles, sciatica, back pain, neuralgia, and headaches including migraines. Infused
oil can be used to treat sore muscles, cold sores and genital herpes, and can be used as a
sunscreen.
• Achillea millefolium—Yarrow. Antibacterial, pain relieving. Herb for wounds. Regulates
blood flow, so that it stops excessive bleeding while also preventing blood from pooling.
Can be used to prevent and treat colds and flu. Diaphoretic. Digestive stimulant.
• Malva spp—Mallow. Whole plant is mucilaginous, extracted in cold water or vinegar, which
is soothing internally (easing sore throats, upset tummies, heart burn, irritable bowel, col-
ic, and constipation) and externally (relieving bug bites, burns, sprains, and sore eyes).
• Impatiens capensis—Jewelweed. Often grows near poison ivy and can be used as an anti-
dote (for poison oak as well)

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Types of Weed 47

• Verbascum spp—Mullein. The leaves are an expectorant, stimulating coughing to clear the
lungs. An infused oil can be used for ear infections. All parts are stimulating to the lymphat-
ic system and are helpful for any ailment related to the alignment of joint, bone or tissue.
• Bellis perennis—Daisy. Relieves headaches, muscle pain and allergy symptoms. Different
from the native daisy (Lagenifera petiolata).
• Ambrosia trifida—Giant ragweed. A tincture can be made and used in small doses to treat
ragweed (and other) allergies.

Other
• Nightshade—breaks up hardpan, allowing roots to grow deeper

Allium Vineale
Allium vineale (wild garlic, crow garlic or stag’s garlic) is a perennial, bulb-forming species of wild
onions, native to Europe, northwestern Africa and the Middle East. The species was introduced in
Australia and North America, where it has become a noxious weed.

Description
All parts of the plant have a strong garlic odour. The underground bulb is 1-2 cm diameter, with
a fibrous outer layer. The main stem grows to 30-120 cm tall, bearing 2-4 leaves and an apical
inflorescence 2-5 cm diameter comprising a number of small bulbils and none to a few flowers,
subtended by a basal bract. The leaves are slender hollow tubes, 15-60 cm long and 2-4 mm thick,
waxy texture, with a groove along the side of the leaf facing the stem. The inflorescence is a tight
umbel surrounded by a membranous bract in bud which withers when the flowers open. Each
individual flower is stalked and has a pinkish-green perianth 2.5 to 4.5 mm (0.10 to 0.18 in) long.
There are six tepals, six stamens and a pistil formed from three fused carpels. Mixed with the
flowers are several of yellowish-brown bulbils. The fruit is a capsule but the seeds seldom set and
propagation usually takes place when the bulbils are knocked off and grow into new plants. Plants
with no flowers, only bulbils, are sometimes distinguished as the variety Allium vineale var. com-
pactum, but this character is probably not taxonomically significant.

Uses and Problems


While Allium vineale has been suggested as a substitute for garlic, there is some difference of
opinion as to whether there is an unpleasant aftertaste compared to that of common garlic (A.
sativum). It imparts a garlic-like flavour and odour on dairy and beef products when grazed by
livestock. It is considered a pestilential invasive weed, as grain products may become tainted with
a garlic odour or flavour in the presence of aerial bulblets at the time of harvest. Wild garlic is resis-
tant to herbicides, which cannot cling well to the vertical, smooth and waxy structure of its leaves.

Arctium Lappa
Arctium lappa, commonly called greater burdock, gobō, edible burdock, lappa, beggar’s buttons,
thorny burr, or happy major is a Eurasian species of plants in the sunflower family, cultivated in

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48 Fundamentals of Weed Science

gardens for its root used as a vegetable. It has become an invasive weed of high-nitrogen soils in
North America, Australia, and other regions.

Arctium lappa - MHNT

Description

Inflorescence.

Greater Burdock is a biennial plant, rather tall, reaching as much as 3 m (10 ft). It has large, alter-
nating, cordiform leaves that have a long petiole and are pubescent on the underside.

The flowers are purple and grouped in globular capitula, united in clusters. They appear in
mid-summer, from July to September. The capitula are surrounded by an involucre made out of
many bracts, each curving to form a hook, allowing them to be carried long distances on the fur of
animals. The fruits are achenes; they are long, compressed, with short pappuses. The fleshy tap-
root can grow up to 1 m (3 ft 3 in) deep.

Distribution and Ecology


This species is native to the temperate regions of the old world, from Scandinavia to the Mediter-
ranean, and from the British Isles through Russia, and the Middle East to India, China, Taiwan
and Japan.

It is naturalized almost everywhere and is usually found in disturbed areas, especially in soil rich
in nitrogen. It is commonly cultivated in Japan where it gives its name to a particular construction
technique, burdock piling.

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Types of Weed 49

The leaves of Greater Burdock provide food for the caterpillars of some Lepidoptera, such as the
Thistle Ermine (Myelois circumvoluta).

Cultivation
It prefers a fresh, worked soil, rich in humus, and should be positioned in full sunlight. Burdock
is very reactive to nitrogen fertilizer. Propagation is achieved through sowing the seeds midsum-
mer. The harvest occurs three to four months after the seeding until late autumn, when the roots
become too fibrous.

Culinary Use

A Japanese appetizer, kinpira gobō, consisting of sauteed gobō (Greater burdock root) and carrot, with a side of
sautéed dried daikon

“Gobō salad” Japanese burdock salad

Greater burdock root is known as niúbàng (牛蒡) in Chinese, which was borrowed into Japanese
as gobō and Korean as ueong (우엉), and is widely eaten in the region. It was used in Europe
during the Middle Ages as a vegetable, but now it is rarely used except in Italy, Brazil and Portugal,
where it is known as bardana or “garduna”. Plants are cultivated for their slender roots, which can
grow about 1 meter long and 2 cm across. The root was traditionally used in Britain as a flavouring
in the herbal drink dandelion and burdock, which is still commercially produced.

Immature flower stalks may also be harvested in late spring, before flowers appear. The taste re-
sembles that of artichoke, to which the burdock is related.

In the second half of the 20th century, burdock achieved international recognition for its culinary
use due to the increasing popularity of the macrobiotic diet, which advocates its consumption. The

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50 Fundamentals of Weed Science

root contains a fair amount of dietary fiber (GDF, 6g per 100g), calcium, potassium, amino acids,
and is low calorie. It contains polyphenols that causes darkened surface and muddy harshness by
formation of tannin-iron complexes. Those polyphenols are caffeoylquinic acid derivatives.

The root is very crisp and has a sweet, mild, and pungent flavor with a little muddy harshness
that can be reduced by soaking julienned/shredded roots in water for five to ten minutes. The
harshness shows excellent harmonization with pork in miso soup (tonjiru) and takikomi gohan (a
Japanese-style pilaf).

A popular Japanese dish is kinpira gobō, julienned or shredded burdock root and carrot, braised
with soy sauce, sugar, mirin and/or sake, and sesame oil. Another is burdock makizushi, rolled
sushi filled with pickled burdock root; the burdock root is often artificially colored orange to re-
semble a carrot. Burdock root can also be found as a fried snack food similar in taste and texture
to potato chips and is occasionally used as an ingredient in tempura dishes.

Fermentation of the root by Aspergillus oryzae is also used for making miso and rice wine in Jap-
anese cuisine.

Use in Traditional Medicine


Dried burdock roots (Bardanae radix) are used in folk medicine as a diuretic, diaphoretic, and
a blood purifying agent. Anecdotal reports from the 19th century suggest that this medicinal
plant has also been used by the Ojibwa tribe, and today, in form of an ingredient in Essiac tea
for the alternative treatment of some cancers. As an oily macerate, it is a component of some
cosmetics, shampoos and hair care products. The seeds of greater burdock are employed in tra-
ditional Chinese medicine particularly for skin conditions and in cold/flu formulas, under the
name niubangzi.

Chemical Constituents
Burdock roots contain mucilage, sulfurous acetylene compounds, polyacetylenes and bitter guaian-
olide-type constituents. Seeds contain arctigenin, arctiin, and butyrolactone lignans.

Mimosa Pudica
Mimosa pudica (from pudica “shy, bashful or shrinking”; also called sensitive plant, sleepy plant,
Dormilones or shy plant) is a creeping annual or perennial herb of the pea family Fabaceae often
grown for its curiosity value: the compound leaves fold inward and droop when touched or
shaken, defending themselves from harm, and re-open a few minutes later. The species is native
to South America and Central America, but is now a pantropical weed. It can also be found in Asia
in countries such as Bangladesh, Thailand, India, Indonesia, Malaysia, Philippines, Japan, and
Jamaica. It grows mostly in undisturbed shady areas, under trees or shrubs.

Taxonomy
Mimosa pudica was first formally described by Carl Linnaeus in Species Plantarum in 1753. The spe-
cies epithet, pudica, is Latin for “bashful” or “shrinking”, alluding to its shrinking reaction to contact.

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Types of Weed 51

The species is known by numerous common names including sensitive plant, humble plant,
shameplant, touch-me-not, muttidare-muni, nachike mullu in kannada, chuimui in Hindi and
Urdu, lajalu in Marathi, lajuki lata in Assamese, Thottanchinungi in Tamil,tottalvaadi in ma-
layalam.

Description

Mimosa pudica flower from Thrissur, Kerala, India

Flower

Mimosa pudica folding leaflets inward.

The whole plant of Mimosa pudica includes thorny stem and branches, flower head, dry flowers,
seed pods, and folded and unfolded leaflets

The stem is erect in young plants, but becomes creeping or trailing with age. It can hang very low
and become floppy. The stem is slender, branching, and sparsely to densely prickly, growing to a
length of 1.5 m (5 ft).

The leaves are bipinnately compound, with one or two pinnae pairs, and 10–26 leaflets per
pinna. The petioles are also prickly. Pedunculate (stalked) pale pink or purple flower heads
arise from the leaf axils in mid summer with more and more flowers as the plant gets older.
The globose to ovoid heads are 8–10 mm in diameter (excluding the stamens). On close exam-
ination, it is seen that the floret petals are red in their upper part and the filaments are pink to
lavender. The fruit consists of clusters of 2–8 pods from 1–2 cm long each, these being prickly

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52 Fundamentals of Weed Science

on the margins. The pods break into 2–5 segments and contain pale brown seeds some 2.5 mm
long. The flowers are pollinated by the wind and insects. The seeds have hard seed coats which
restrict germination.

The roots of Mimosa pudica create carbon disulfide, which prevents certain pathogenic and my-
corrhizal fungi from growing within the plant’s rhizosphere. This allows the formation of nodules
on the roots of the plant that contain endosymbiotic diazotrophs, which fix atmospheric nitrogen
and convert it into a form that is usable by the plant.

Mimosa pudica is a tetraploid (2n = 52).

Plant Movement

Video of Mimosa pudica closing when touched in Hainan, China

Mimosa pudica is well known for its rapid plant movement. Like a number of other plant spe-
cies, it undergoes changes in leaf orientation termed “sleep” or nyctinastic movement. The foliage
closes during darkness and reopens in light. This was first studied by the French scientist Jean-
Jacques d’Ortous de Mairan.

The leaves also close under various other stimuli, such as touching, warming, blowing, or shaking.
These types of movements have been termed seismonastic movements. The stimulus is transmit-
ted via action potentials to the pulvini of the plant, where contractile proteins and aquaporins
induce the movement of water and electrolytes out of the cell. The movement occurs when specific
regions of cells lose turgor pressure, which is the force that is applied onto the cell wall by water
within the cell vacuoles and other cell contents. When the plant is disturbed, specific regions on
the stems are stimulated to release chemicals, including potassium ions, which force water out
of the cell vacuoles and the water diffuses out of the cells, producing a loss of cell pressure and
cell collapse; this differential turgidity between different regions of cells results in the closing of
the leaflets and the collapse of the leaf petiole. This movement of folding inwards is energetically
costly for the plant and also interferes with the process of photosynthesis. This characteristic is
quite common within the Mimosoideae subfamily of the legume family, Fabaceae. The stimulus
can also be transmitted to neighboring leaves. It is not known exactly why Mimosa pudica evolved
this trait, but many scientists think that the plant uses its ability to shrink as a defense from herbi-
vores. Animals may be afraid of a fast moving plant and would rather eat a less active one. Another
possible explanation is that the sudden movement dislodges harmful insects.

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Types of Weed 53

Distribution and Habitat


Mimosa pudica is native to South America, North America and Central America. It has been in-
troduced to many other regions and is regarded as an invasive species in Tanzania, South Asia and
South East Asia and many Pacific Islands. It is regarded as invasive in parts of Australia and is a
declared weed in the Northern Territory, and Western Australia although not naturalized there.
Control is recommended in Queensland. It has also been introduced to Ghana, Nigeria, Seychelles,
Mauritius and East Asia but is not regarded as invasive in those places. In the United States of
America, it grows in Florida, Hawaii, Tennessee, Virginia, Maryland, Puerto Rico, Texas, Alabama,
Mississippi, North Carolina, Georgia, and the Virgin Islands, as well as Cuba and the Dominican
Republic.

Agricultural Impacts
The species can be a troublesome weed in tropical crops, particularly when fields are hand cultivat-
ed. Crops it tends to affect are corn, coconuts, tomatoes, cotton, coffee, bananas, soybeans, papaya,
and sugar cane. Dry thickets may become a fire hazard. In some cases it has become a forage plant
although the variety in Hawaii is reported to be toxic to livestock.

In addition, Mimosa pudica can change the physico-chemical properties of the soil it invades. For
example, the total N and K increased in significantly invaded areas.

Nitrogen Fixation
Mimosa pudica can form root nodules that are habitable by nitrogen-fixing bacteria. The bacteria
are able to convert atmospheric nitrogen, which plants cannot use, into a form that plants can use.
This trait is common among plants in the Fabaceae family. Nitrogen is a vital element for both
plant growth and reproduction. Nitrogen is also essential for plant photosynthesis because it is a
component of chlorophyll. Nitrogen fixation contributes nitrogen to the plant and to the soil sur-
rounding the plant’s roots.

Mimosa pudica’s ability to fix nitrogen may have arisen in conjunction with the evolution of ni-
trogen-fixing bacteria. Nitrogen fixation is an adaptive trait that has transformed the parasitic
relationship between the bacteria and plants into a mutualistic relationship. The shifting dynam-
ics of this relationship are demonstrated by the corresponding improvement of various symbiotic
characteristics in both Mimosa pudica and bacteria. These traits include enhanced “competitive
nodulation, nodule development, intracellular infection, and bacteroid persistence”. As much as
60% of the nitrogen found in Mimosa pudica can be attributed to the fixation of N2 by bacteria.
Burkholderia phymatum STM815T and Cupriavidus taiwanensis LMG19424T are beta-rhizobial
strains of diazotrophs that are highly effective at fixing nitrogen when coupled with M. pudica.
Burkholderia is also shown to be a strong symbiont of Mimosa pudica in nitrogen-poor soils in
regions like Cerrado and Caatinga.

Cultivation
In cultivation, this plant is most often grown as an indoor annual, but is also grown for groundcov-
er. Propagation is generally by seed. Mimosa pudica grows most effectively in nutrient poor soil

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54 Fundamentals of Weed Science

that allows for substantial water drainage. However, this plant is also shown to grow in scalped
and eroded subsoils. Typically, disrupted soil is necessary in order for M. pudica to become es-
tablished in an area. Additionally, the plant is shade intolerant and frost-sensitive, meaning that
it does not tolerate low levels of light or cold temperatures. Mimosa pudica does not compete for
resources with larger foliage or forest canopy undergrowth.

Chemical Constituents
Mimosa pudica contains the toxic alkaloid mimosine, which has been found to also have an-
tiproliferative and apoptotic effects. The extracts of Mimosa pudica immobilize the filariform
larvae of Strongyloides stercoralis in less than one hour. Aqueous extracts of the roots of the
plant have shown significant neutralizing effects in the lethality of the venom of the monocled
cobra (Naja kaouthia). It appears to inhibit the myotoxicity and enzyme activity of cobra ven-
om.

Mimosa pudica demonstrates both antioxidant and antibacterial properties. This plant has also
been demonstrated to be non-toxic in brine shrimp lethality tests, which suggests that M. pudica
has low levels of toxicity. Chemical analysis has shown that Mimosa pudica contains various com-
pounds, including “alkaloids, flavonoid C-glycosides, sterols, terenoids, tannins, and fatty acids”.
The roots of the plant have been shown to contain up to 10% tannin. A substance similar to adren-
aline has been found within the plant’s leaves. Mimosa pudica’s seeds produce mucilage made up
of D-glucuronic acid and D-xylose. Additionally, extracts of M. pudica have been shown to contain
crocetin-dimethylester, tubulin, and green-yellow fatty oils. A new class of phytohormone turgo-
rines, which are derivatives of gallic acid 4-O-(β-D-glucopyranosyl-6’-sulfate), have been discov-
ered within the plant.

The nitrogen-fixing properties of Mimosa pudica contribute to a high nitrogen content with-
in the plant’s leaves. The leaves of M. pudica also contain a wide range of carbon to mineral
content, as well as, a large variation in 13C values. The correlation between these two numbers
suggests that significant ecological adaptation has occurred among the varieties of M. pudica
in Brazil.

The roots contain sac-like structures that release organic and organosulfur compounds includ-
ing SO2, methylsulfinic acid, pyruvic acid, lactic acid, ethanesulfinic acid, propane sulfinic acid,
2-mercaptoaniline, S-propyl propane 1-thiosulfinate, and thioformaldehyde, an elusive and highly
unstable compound never before reported to be emitted by a plant.

Rumex

The docks and sorrels, genus Rumex L., are a genus of about 200 species of annual, biennial, and
perennial herbs in the buckwheat family Polygonaceae.

Members of this family are very common perennial herbs growing mainly in the Northern Hemi-
sphere, but various species have been introduced almost everywhere.

Some are nuisance weeds (and are sometimes called dockweed or dock weed), but some are grown
for their edible leaves.

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Rumex species are used as food plants by the larvae of a number of Lepidoptera species.

Description

They are erect plants, usually with long taproots. The fleshy to leathery leaves form a basal rosette
at the root. The basal leaves may be different from those near the inflorescence. They may or may
not have stipules. Minor leaf veins occur. The leaf blade margins are entire or crenate.

The usually inconspicuous flowers are carried above the leaves in clusters. The fertile flowers are
mostly hermaphrodites, or they may be functionally male or female. The flowers and seeds grow on
long clusters at the top of a stalk emerging from the basal rosette; in many species, the flowers are
green, but in some (such as sheep’s sorrel, Rumex acetosella) the flowers and their stems may be
brick-red. Each seed is a three-sided achene, often with a round tubercle on one or all three sides.

Uses

These plants have many uses. Broad-leaved dock (Rumex obtusifolius) used to be called butter
dock because its large leaves were used to wrap and conserve butter.

Rumex hymenosepalus has been cultivated in the Southwestern US as a source of tannin (roots
contain up to 25%), for use in leather tanning, while leaves and stems are used for a mordant-free
mustard-colored dye.

These plants are edible. The leaves of most species contain oxalic acid and tannin, and many have
astringent and slightly purgative qualities. Some species with particularly high levels of oxalic acid
are called sorrels (including sheep’s sorrel, Rumex acetosella, common sorrel, Rumex acetosa, and
French sorrel, Rumex scutatus), and some of these are grown as leaf vegetables or garden herbs
for their acidic taste.

In Western Europe, dock leaves are a traditional remedy for the sting of nettles, and suitable larger
docks (such as broad-leaved dock R. obtusifolius or curled dock R. crispus) often grow convenient-
ly in similar habitats to the common nettle (Urtica dioica).

In traditional Austrian medicine, R. alpinus leaves and roots have been used internally for treat-
ment of viral infections.

Species

Broad-leaved dock leaves (R. obtusifolius)

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56 Fundamentals of Weed Science

Flowers of curled dock (R. crispus) with remarkable tubercles

• Rumex acetosa – sorrel, common sorrel, garden sorrel, narrow-leaved dock, spinach dock

• Rumex acetosella - sheep’s sorrel, common sheep sorrel, field sorrel, red sorrel

o Rumex acetosella ssp. acetosella

o Rumex acetosella ssp. acetoselloides

o Rumex acetosella ssp. multifidus

o Rumex acetosella ssp. pyrenaicus - round-seeded sheep’s sorrel

• Rumex x acutus - acute dock

• Rumex albescens - Oahu dock

• Rumex x alexidis - Alex’s dock

• Rumex arifolius - maiden sorrel, mountain dock

• Rumex alpinus - alpine dock, monk’s rhubarb

• Rumex altissimus - pale dock, smooth dock, peach-leaf dock

• Rumex andinus

• Rumex angiocarpus

• Rumex aquaticus L. – western dock

o Rumex aquaticus ssp. aquaticus

o Rumex aquaticus ssp. arcticus - arctic dock

o Rumex aquaticus ssp. protractus

• Rumex aquaticus x longifolius

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Types of Weed 57

• Rumex aquitanicus

• Rumex arcticus

• Rumex azoricus

• Rumex balcanicus

• Rumex beringensis - Bering Sea dock

• Rumex brownei - Browne’s dock

• Rumex brownii - swamp dock

• Rumex bucephalophorus - red dock

o Rumex bucephalophorus subsp. bucephalophorus

o Rumex bucephalophorus subsp. aegaeus

o Rumex bucephalophorus subsp. canariensis

o Rumex bucephalophorus subsp. gallicus

o Rumex bucephalophorus subsp. hispanicus

o Rumex bucephalophorus var. aegaeus

o Rumex bucephalophorus var. subaegaeus

• Rumex chrysocarpus - amamastla

• Rumex confertus Willd. – Asiatic Dock

• Rumex x confusus - confused dock

• Rumex conglomeratus – clustered dock, sharp dock

• Rumex costaricensis

• Rumex crispus – curled dock, curly dock, yellow dock, sour dock, narrow dock, garden
patience, narrow-leaved dock

o Rumex crispus subsp. crispus

o Rumex crispus subsp. littoreus

o Rumex crispus subsp. uliginosus

• Rumex crispus x confertus

• Rumex crispus x longifolius

• Rumex crispus x obtusifolius

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58 Fundamentals of Weed Science

• Rumex crispus x patientia

• Rumex crispus x pulcher

• Rumex cristatus : Greek Dock

o Rumex cristatus subsp. cristatus

o Rumex cristatus subsp. kerneri

• Rumex crystallinus - Shiny Dock

• Rumex densiflorus - dense-flower or -flowered dock

• Rumex dentatus - toothed dock

o Rumex dentatus subsp. dentatus

o Rumex dentatus subsp. halacsyi

o Rumex dentatus subsp. reticulatus

• Rumex diclinis

• Rumex digynus

• Rumex dimidiatus

• Rumex dimorphophyllus

• Rumex x dissimilis - dock

• Rumex x dobrogensis

• Rumex x dolosus

• Rumex dregeanus

• Rumex drobovii

• Rumex drummondii

• Rumex × dufftii

• Rumex dumosiformis

• Rumex dumosus - wiry dock

o Rumex dumosus var. dumosus

• Rumex × dumulosus

• Rumex durispissimus

• Rumex ecklonianus

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• Rumex ecuadoriensis

• Rumex elbrusensis

• Rumex ellenbeckii

• Rumex engelmanni

• Rumex ephedroides

• Rumex erosus

• Rumex x erubescens

• Rumex erythrocarpus

• Rumex esquirolii

• Rumex euxinus

• Rumex evenkiensis

• Rumex x exspectatus

• Rumex x fallacinus

• Rumex fascicularis

• Rumex fascilobus

o Rumex fenestratus subsp. puberulus

• Rumex fimbriatus

• Rumex x finitimus

• Rumex flexicaulis

• Rumex flexuosiformis

• Rumex flexuosus

• Rumex foliosus

• Rumex fontano-paludosus

• Rumex foveolatus

• Rumex x franktonis - Frankton dock

• Rumex fraternus

• Rumex fringillimontanus

• Rumex frutescens - wedgeleaf dock

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60 Fundamentals of Weed Science

• Rumex fueginus

o Rumex fueginus var. brachythrix

o Rumex fueginus var. ovato-cordatus

o Rumex fueginus var. tanythrix

o Rumex fueginus var. typicus

• Rumex gamsii

• Rumex gangotrianus

• Rumex gieshueblensis

• Rumex giganteus - Pawale

• Rumex ginii

• Rumex gmelini

• Rumex gmelinii var. latus

• Rumex gombae

• Rumex gracilescens

• Rumex gracilipes

• Rumex graminifolius - grassleaf sorrel

• Rumex granulosus

• Rumex x griffithii

• Rumex x grintzescui

• Rumex gussonii

• Rumex x gusuleacii

• Rumex hadmocarpus

• Rumex halophilus

• Rumex hararensis

• Rumex hasslerianus

• Rumex hastatulus - heartwing dock, heartwing sorrel

• Rumex hastatus

• Rumex hayekii

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Types of Weed 61

• Rumex hazslinszkyanus

• Rumex x heimerlii

• Rumex hellenicus

• Rumex henrardi

• Rumex hesperius

• Rumex × heteranthos

• Rumex heterophylus

• Rumex hexagynus

• Rumex hippiatricus

• Rumex hirsutus

• Rumex horizontalis

• Rumex hoschedei

• Rumex hostilis

• Rumex hultenii

• Rumex hungaricus

• Rumex x hybridus

• Rumex hydrolapathum - great water dock

• Rumex hymenosepalus - canaigre, canaigre dock

• Rumex x impurus

• Rumex inconspicuus

• Rumex integer

• Rumex integrifolia

• Rumex x intercedens

• Rumex intermedius

o Rumex intermedius ssp. algarbiensis

o Rumex intermedius ssp. lusitanicus

• Rumex interruptus

• Rumex x inundatus

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62 Fundamentals of Weed Science

• Rumex iseriensis

• Rumex jacutensis

• Rumex japonicus

• Rumex x johannis-moorei

• Rumex kamtshadalus

• Rumex kaschgaricus

• Rumex x kaschmirianus

• Rumex kerneri - Kerner’s dock

• Rumex khekii

• Rumex x khorasanicus

• Rumex x knafii

• Rumex komarovii

• Rumex krausei - Krause’s sorrel

• Rumex lachanus

• Rumex lacustris

• Rumex lanceolatus

• Rumex langloisii

• Rumex lanuginosus

• Rumex lapponicus

• Rumex lanuginosus

• Rumex latifolius

• Rumex lativalvis

• Rumex leptocaulis

• Rumex leptophyllus

• Rumex limoniastrum

• Rumex linearis

• Rumex x lingulatus

• Rumex litoralis

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Types of Weed 63

• Rumex lonaczewskii

• Rumex longifolius – dooryard dock, northern dock

• Rumex longisetus

• Rumex lorentzianus

• Rumex x lousleyi

• Rumex ludovicianus

• Rumex lugdunensis

• Rumex lunaria

• Rumex luxurians

• Rumex x lycheanus

• Rumex maderensis

• Rumex magellanicus

o Rumex magellanicus var. donatii

o Rumex magellanicus var. dusenii

• Rumex maritimus – golden dock, bristle dock, seashore dock

• Rumex marschallianus

• Rumex maximus

• Rumex megalophyllus

• Rumex meyeri

• Rumex × mezei

• Rumex microcarpus

• Rumex microdon

• Rumex x mirabilis

• Rumex mixtus

• Rumex moedlingensis

• Rumex x monistrolensis

• Rumex montanus

• Rumex monticola

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64 Fundamentals of Weed Science

• Rumex muelleri

• Rumex x munshii

• Rumex muretii

• Rumex muricatus

• Rumex x nankingensis

• Rumex natalensis

• Rumex neglectus

• Rumex nematopodus - Arizona dock

• Rumex nemorosus

• Rumex nepalensis

• Rumex nervosus

• Rumex nevadensis

• Rumex nigricans

• Rumex nikkoensis

• Rumex nivalis

• Rumex oblongifolius

• Rumex obovatus Danser – tropical dock

• Rumex obtusifolius – broad-leaved dock, bitter dock, bluntleaf dock, butter dock

• Rumex obtusifolius ssp. obtusifolius

• Rumex occidentalis

o Rumex occidentalis var. perplexus

• Rumex occultans

• Rumex ochotensis

• Rumex orbiculatus - great water dock

• Rumex orientalis

• Rumex orthoneurus - Chiricahua mountain dock

• Rumex x oryzetorum

• Rumex osswaldii

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Types of Weed 65

• Rumex oxysepalus

• Rumex x pakistanicus

• Rumex pallidus - seaside dock

• Rumex palustris – marsh dock

• Rumex x palustroides

• Rumex pamiricus

• Rumex x pannonicus

• Rumex papilio

• Rumex paraguayensis - Paraguayan dock

• Rumex patagonicus

• Rumex patientia – patience dock, garden patience, monk’s rhubarb

• Rumex paucifolius - alpine sheep’s sorrel, few-leaved dock, meadow dock

• Rumex peisonis

• Rumex peregrinus

• Rumex persicaris

• Rumex peruanus

• Rumex pictus

• Rumex pilatensis

• Rumex planivalvis

• Rumex polycarpus

• Rumex polygamus

• Rumex polyklonos

• Rumex polyrrhizus

• Rumex ponticus

• Rumex popovii

• Rumex praecox

• Rumex pratensis

• Rumex procerus

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66 Fundamentals of Weed Science

• Rumex x promiscuus

• Rumex x propinquus

• Rumex protractus

• Rumex pseudonatronatus - field dock

• Rumex x pseudopulcher

• Rumex pseudoscutatus

• Rumex pseudoxyria

• Rumex pulcher - fiddle dock

o Rumex pulcher ssp. divaricatus

o Rumex pulcher ssp. pulcher

o Rumex pulcher ssp. woodsii

• Rumex quarrei

• Rumex raulini

• Rumex rechingerianus

• Rumex rectinervius

• Rumex recurvatus

• Rumex x rhaeticus

• Rumex rhodesius

• Rumex x romanicus

• Rumex romassa

• Rumex x rosemurphyae

• Rumex roseus

• Rumex rossicus

• Rumex rothschildianus

• Rumex rupestris - Shore Dock

• Rumex ruwenzoriensis

• Rumex sagittatus - rambling dock

• Rumex x sagorski

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Types of Weed 67

• Rumex salicetorum

• Rumex salicifolius - willow dock, willow-leaved dock

o Rumex salicifolius ssp. angustivalvis

o Rumex salicifolius var. denticulatus

o Rumex salicifolius f. escallosus

o Rumex salicifolius ssp. montigenitus

o Rumex salicifolius var. nudivalvis

o Rumex salicifolius var. oreolapathum - willow dock

o Rumex salicifolius var. triangularis

o Rumex salicifolius var. trigranis

• Rumex salinus

• Rumex samuelssoni

• Rumex sanguineus L. – wood dock, redvein dock

o Rumex sanguineus var. sanguineus - bloody dock, blood dock, bloodwort, bloody
wood dock

• Rumex sanninensis

• Rumex schimperi

• Rumex schischkinii

• Rumex x schmidtii

• Rumex x schreberi

• Rumex x schultzei

• Rumex scutatus - French sorrel, leaf-shield sorrel

o Rumex scutatus ssp. gallaecicus

• Rumex sellowianus

• Rumex semigraecus

• Rumex shultzii

• Rumex sibiricus - Siberian dock

• Rumex similans

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68 Fundamentals of Weed Science

• Rumex x similatus

• Rumex simonkaianus

• Rumex simpliciflorus

• Rumex sinuatus

• Rumex x skofitzi

• Rumex skottsbergii - lava dock

• Rumex songaricus

• Rumex x sorkhabadensis

• Rumex spathulatus

• Rumex spiralis - winged dock

• Rumex spurius

• Rumex stenoglottis

• Rumex stenophyllus

• Rumex subalpinus - subalpine dock

• Rumex subarcticus

• Rumex x subdubius

• Rumex subrotundus

• Rumex subtrilobus

• Rumex subvirescens

• Rumex suffruticosus

• Rumex suzukianus

• Rumex sylvaticus

• Rumex sylvestris

• Rumex x talaricus

• Rumex tenax

• Rumex tenellus

• Rumex thyrsiflorus

• Rumex thyrsoides

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Types of Weed 69

• Rumex tianschanicus

• Rumex tibeticus

• Rumex tingitanus

• Rumex tmoleus

• Rumex toepffieri

• Rumex tolimensis

• Rumex tomentellus - willow creek dock

• Rumex × transbaicalicus

• Rumex triangularis

• Rumex triangulivalvis

• Rumex trifidus

• Rumex trinervius

• Rumex tuberosus

• Rumex tunetanus

• Rumex turcestanicus

• Rumex ucranicus

• Rumex x ujskensis

• Rumex x uludaghensis

• Rumex uncinulatus

• Rumex undulatus

• Rumex ursinus

• Rumex uruguayensis

• Rumex ussuriensis

• Rumex usticanus

• Rumex utahensis

• Rumex venosus - veiny dock or sand dock

• Rumex verrietianus

• Rumex verticillatus - swamp dock, water dock

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70 Fundamentals of Weed Science

• Rumex vesceritensis

• Rumex vesicarius - bladder dock

• Rumex violascens - violet dock

• Rumex wachteri

• Rumex x weberi

• Rumex × wildtianus

• Rumex woodii

• Rumex woodsii

• Rumex x wrightii

• Rumex × xenogenus

• Rumex yezoensis

• Rumex yungningensis

• Rumex zsakii

Blackberry
The blackberry is an edible fruit produced by many species in the Rubus genus in the Rosaceae
family, hybrids among these species within the Rubus subgenus, and hybrids between the Rubus
and Idaeobatus subgenera. The taxonomy of the blackberries has historically been confused be-
cause of hybridization and apomixis, so that species have often been grouped together and called
species aggregates. For example, the entire subgenus Rubus has been called the Rubus fruticosus
aggregate, although the species R. fruticosus is considered a synonym of R. plicatus.

Description
What distinguishes the blackberry from its raspberry relatives is whether or not the torus (receptacle or
stem) “picks-with” (i.e. stays with) the fruit. When picking a blackberry fruit, the torus does stay with
the fruit. With a raspberry, the torus remains on the plant, leaving a hollow core in the raspberry fruit.

The term “bramble”, a word meaning any impenetrable thicket, has traditionally been applied
specifically to the blackberry or its products, though in the United States it applies to all members
of the Rubus genus. In the western US, the term caneberry is used to refer to blackberries and
raspberries as a group rather than the term bramble.

The usually black fruit is not a berry in the botanical sense of the word. Botanically it is termed
an aggregate fruit, composed of small drupelets. It is a widespread and well-known group of over
375 species, many of which are closely related apomictic microspecies native throughout Europe,
northwestern Africa, temperate western and central Asia and North and South America.

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Types of Weed 71

Botanical Characteristics
Blackberries are perennial plants which typically bear biennial stems (“canes”) from the perennial
root system.

In its first year, a new stem, the primocane, grows vigorously to its full length of 3–6 m (in some
cases, up to 9 m), arching or trailing along the ground and bearing large palmately compound
leaves with five or seven leaflets; it does not produce any flowers. In its second year, the cane
becomes a floricane and the stem does not grow longer, but the lateral buds break to produce
flowering laterals (which have smaller leaves with three or five leaflets). First- and second-year
shoots usually have numerous short-curved, very sharp prickles that are often erroneously called
thorns. These prickles can tear through denim with ease and make the plant very difficult to nav-
igate around. Prickle-free cultivars have been developed. Recently the University of Arkansas has
developed primocane fruiting blackberries that grow and flower on first-year growth much as the
primocane-fruiting (also called fall bearing or everbearing) red raspberries do.

Unmanaged mature plants form a tangle of dense arching stems, the branches rooting from the
node tip on many species when they reach the ground. Vigorous and growing rapidly in woods,
scrub, hillsides, and hedgerows, blackberry shrubs tolerate poor soils, readily colonizing waste-
land, ditches, and vacant lots.

The flowers are produced in late spring and early summer on short racemes on the tips of the flow-
ering laterals. Each flower is about 2–3 cm in diameter with five white or pale pink petals.

The drupelets only develop around ovules that are fertilized by the male gamete from a pollen
grain. The most likely cause of undeveloped ovules is inadequate pollinator visits. Even a small
change in conditions, such as a rainy day or a day too hot for bees to work after early morning, can
reduce the number of bee visits to the flower, thus reducing the quality of the fruit. Incomplete
drupelet development can also be a symptom of exhausted reserves in the plant’s roots or infection
with a virus such as Raspberry bushy dwarf virus.

Ecology
Blackberry leaves are food for certain caterpillars; some grazing mammals, especially deer, are also
very fond of the leaves. Caterpillars of the concealer moth Alabonia geoffrella have been found feed-
ing inside dead blackberry shoots. When mature, the berries are eaten and their seeds dispersed by
several mammals, such as the red fox and the Eurasian badger, as well as by small birds.

A bee, Bombus hypnorum, pollinating blackberries

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72 Fundamentals of Weed Science

A basket of wild blackberries

Blackberry fruit are red before they are ripe, leading to an old expression that “blackberries are red
when they’re green”.

In various parts of the United States, wild blackberries are sometimes called “Black-caps”, a term
more commonly used for black raspberries, Rubus occidentalis.

As there is forensic evidence from the Iron Age Haraldskær Woman that she consumed black-
berries some 2500 years ago, it is reasonable to conclude that blackberries have been eaten by
humans over thousands of years.

Uses
Nutrients
American cultivated blackberries (R. laciniatus and R. ursinus) are notable for their significant
contents of dietary fiber, vitamin C, and vitamin K (table). A 100 gram serving of raw blackberries
supplies 43 calories and 5 grams of dietary fiber or 25% of the recommended Daily Value (DV)
(table). In 100 grams, vitamin C and vitamin K contents are 25% and 19% DV, respectively, while
other essential nutrients are low in content (table).

Blackberries contain both soluble and insoluble fiber components.

Nutrient Content of Seeds


Blackberries contain numerous large seeds that are not always preferred by consumers. The seeds
contain oil rich in omega-3 (alpha-linolenic acid) and -6 fats (linoleic acid) as well as protein, di-
etary fiber, carotenoids, ellagitannins and ellagic acid.

Food
The soft fruit is popular for use in desserts, jams, seedless jelly, and sometimes wine. It is often
mixed with apples for pies and crumbles. Blackberries are also used to produce candy.

Good nectar producers, blackberry shrubs bearing flowers yield a medium to dark, fruity honey.

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Types of Weed 73

Wild blackberries picked in May in Texas

Wild Blackberries in Virginia

Phytochemical Research
Blackberries contain numerous phytochemicals including polyphenols, flavonoids, anthocyanins,
salicylic acid, ellagic acid, and fiber. Anthocyanins in blackberries are responsible for their rich
dark color.

Phytochemical components of blackberries, salicylic acid and ellagic acid have been associated in
preliminary research with toxicity to cancer cells, including breast cancer cells.

Blackberries rank highly among fruits for in vitro antioxidant strength, particularly because of
their dense content of polyphenolic compounds, such as ellagic acid, tannins, ellagitannins, quer-
cetin, gallic acid, anthocyanins, and cyanidins. One report placed blackberry at the top of more
than 1000 polyphenol-rich foods consumed in the United States, but this concept of a health bene-
fit from consuming darkly colored foods like blackberries remains scientifically unverified and not
accepted for health claims on food labels.

Cultivation
Commercial Cultivation
Worldwide, Mexico is the leading producer of blackberries, with nearly the entire crop being pro-
duced for export into the off-season fresh markets in North America and Europe. The Mexican
market is almost entirely from the cultivar ‘Tupy’ (often spelled ‘Tupi’, but the EMBRAPA program
in Brazil from which it was released prefers the ‘Tupy’ spelling). In the US, Oregon is the leading
commercial blackberry producer, producing 42.6 million pounds on 6,180 acres (25.0  km2), in
1995 and 56.1 million pounds on 7,000 acres (28 km2) in 2009.

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74 Fundamentals of Weed Science

Numerous cultivars have been selected for commercial and amateur cultivation in Europe and the
United States. Since the many species form hybrids easily, there are numerous cultivars with more
than one species in their ancestry.

‘Marion’ (marketed as “marionberry”) is an important cultivar that was selected from seedlings
from a cross between ‘Chehalem’ and ‘Olallie’ (commonly called “Olallieberry”) berries. ‘Olallie’ in
turn is a cross between loganberry and youngberry. ‘Marion’, ‘Chehalem’ and ‘Olallie’ are just three
of many trailing blackberry cultivars developed by the United States Department of Agriculture
Agricultural Research Service (USDA-ARS) blackberry breeding program at Oregon State Univer-
sity in Corvallis, Oregon.

The most recent cultivars released from this program are the prickle-free cultivars ‘Black Dia-
mond’, ‘Black Pearl’, and ‘Nightfall’ as well as the very early-ripening ‘Obsidian’ and ‘Metolius’.
‘Black Diamond’ is now the leading cultivar being planted in the Pacific Northwest. Some of the
other cultivars from this program are ‘Newberry’, ‘Waldo’, ‘Siskiyou’, ‘Black Butte’, ‘Kotata’, ‘Pacif-
ic’, and ‘Cascade’.

Trailing blackberries are vigorous and crown forming, require a trellis for support, and are less
cold hardy than the erect or semi-erect blackberries. In addition to the United States’s Pacific
Northwest, these types do well in similar climates such as the United Kingdom, New Zealand,
Chile, and the Mediterranean countries.

Semi-erect, prickle-free blackberries were first developed at the John Innes Centre in Norwich,
UK, and subsequently by the USDA-ARS in Beltsville, Maryland. These are crown forming and
very vigorous and need a trellis for support. Cultivars include ‘Black Satin’ ‘Chester Thornless’,
‘Dirksen Thornless’, ‘Hull Thornless’, ‘Loch Maree’, ‘Loch Ness’, ‘Loch Tay’, ‘Merton Thornless’,
‘Smoothstem’, and ‘Triple Crown’. Recently, the cultivar ‘Cacanska Bestrna’ (also called ‘Cacak
Thornless’) has been developed in Serbia and has been planted on many thousands of hectares
there.

The University of Arkansas has developed cultivars of erect blackberries. These types are less
vigorous than the semi-erect types and produce new canes from root initials (therefore they
spread underground like raspberries). There are prickly and prickle-free cultivars from this
program, including ‘Navaho’, ‘Ouachita’, ‘Cherokee’, ‘Apache’, ‘Arapaho’, and ‘Kiowa’. They
are also responsible for developing the primocane fruiting blackberries such as ‘Prime-Jan’
and ‘Prime-Jim’.

In raspberries, these types are called primocane fruiting, fall fruiting, or everbearing. ‘Prime-Jim’
and ‘Prime-Jan’ were released in 2004 by the University of Arkansas and are the first cultivars
of primocane fruiting blackberry. They grow much like the other erect cultivars described above,
however the canes that emerge in the spring, will flower in mid-summer and fruit in late summer
or fall. The fall crop has its highest quality when it ripens in cool mild climate such as in California
or the Pacific Northwest.

‘Illini Hardy’ a semi-erect prickly cultivar introduced by the University of Illinois is cane hardy in
zone 5, where traditionally blackberry production has been problematic, since canes often failed
to survive the winter.

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Types of Weed 75

Blackberry production in Mexico has expanded enormously in the past decade. While once based
on the cultivar ‘Brazos’, an old erect blackberry cultivar developed in Texas in 1959, the Mexi-
can industry is now dominated by the Brazilian ‘Tupy’ released in the 1990s. ‘Tupy’ has the erect
blackberry ‘Comanche’, and a “wild Uruguayan blackberry” as parents. Since there are no native
blackberries in Uruguay, the suspicion is that the widely grown ‘Boysenberry’ is the male parent.
In order to produce these blackberries in regions of Mexico where there is no winter chilling to
stimulate flower bud development, chemical defoliation and application of growth regulators are
used to bring the plants into bloom.

Diseases and Pests

Raindrop on blackberry pale pink flower

As a result of blackberries belonging to the same genus as raspberries, they share the same dis-
eases including anthracnose which can cause the berry to have uneven ripening and sap flow may
also be slowed. They also share the same remedies including the Bordeaux mixture, a combination
of lime, water and copper(II) sulfate. The rows between blackberry plants must be free of weeds,
blackberry suckers and grasses which may lead to pests or diseases. Fruit growers are selective
when planting blackberry bushes as wild blackberries may be infected and gardeners are recom-
mended to purchase only certified disease-free plants.

The spotted-wing drosophila, Drosophila suzukii is a serious pest of blackberries. Unlike its vine-
gar fly relatives which are primarily attracted to rotting or fermented fruit, D. suzukii attacks fresh,
ripe fruit by laying eggs under the soft skin. The larvae hatch and grow in the fruit, destroying the
fruit’s commercial value.

Another pest is Amphorophora rubi, known as the Blackberry Aphid, which not only eats black-
berries but raspberries as well.

Byturus tomentosus (Raspberry beetle), Lampronia corticella (Raspberry Moth) and Anthono-
mus rubi (Strawberry blossom weevil) are also known to infest blackberries.

Folklore
Folklore in the United Kingdom is told that blackberries should not be picked after Old Michael-
mas Day (11 October) as the devil (or a Púca) has made them unfit to eat by stepping, spitting or

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76 Fundamentals of Weed Science

fouling on them. There is some value in this legend as autumn’s wetter and cooler weather often
allows the fruit to become infected by various molds such as Botryotinia which give the fruit an
unpleasant look and may be toxic. According to some traditions, blackberry’s deep purple color
represents Christ’s blood and the crown of thorns was made of brambles, although other thorny
plants, such as Crataegus (hawthorn species) and Euphorbia milii (crown of thorns plant), have
been proposed as the material for the crown.

Centaurea Cyanus
Centaurea cyanus, commonly known as cornflower, is an annual flowering plant in the family As-
teraceae, native to Europe. In the past it often grew as a weed in cornfields (in the broad sense of
the word “corn”, referring to grains, such as wheat, barley, rye, or oats), hence its name. It is now
endangered in its native habitat by agricultural intensification, particularly over-use of herbicides,
destroying its habitat; in the United Kingdom it has declined from 264 sites to just 3 sites in the
last 50 years. In reaction to this, the conservation charity Plantlife named it as one of 101 species it
would actively work to bring ‘Back from the Brink’. It is also, however, through introduction as an
ornamental plant in gardens and a seed contaminant in crop seeds, now naturalised in many other
parts of the world, including North America and parts of Australia.

Description
Cornflower is an annual plant growing to 16-35 inches tall, with grey-green branched stems. The
leaves are lanceolate, 1–4 cm long. The flowers are most commonly an intense blue colour, pro-
duced in flowerheads (capitula) 1.5–3 cm diameter, with a ring of a few large, spreading ray florets
surrounding a central cluster of disc florets. The blue pigment is protocyanin, which in roses is red.
It flowers all summer.

Distribution
Centaurea cyanus is native to temperate Europe, but is widely naturalized outside its native range.
It has been present in the British Isles as an archaeophyte (ancient introduction) since the Iron
Age. In Ireland, in Co.Clare (H9) Centaurea cyanus is recorded in arable fields as very rare and
almost extinct. while in the north-east of Ireland it was abundant before 1930s.

Names
The common names bachelor’s button, bluebottle, boutonniere flower, hurtsickle or cyani flower
are also used.

It may also be referred to as basketflower, though the term also refers to the Plectocephalus group
of Centaurea, which is probably a distinct genus. “Cornflower” is also used for chicory, and a few
other Centaurea species; to distinguish C. cyanus from these it is sometimes called common corn-
flower.

Cultivation
It is grown as an ornamental plant in gardens, where several cultivars have been selected with

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Types of Weed 77

varying pastel colours, including pink and purple. Cornflower is also grown for the cutflower in-
dustry in Canada for use by florists. The most common colour variety for this use is a doubled blue
variety such as ‘Blue Boy’ or ‘Blue Diadem’. White, pink, lavender and black (actually a very dark
maroon) are also used but less commonly. A relative, Centaurea montana, is a perennial plant
which is also cultivated as a garden plant. Cornflowers germinate quickly after planting.

Dried Cornflower as used in herbal tea & tea blends

Light requirements: full sun. Water requirements: high-average water daily. Soil pH requirements:
neutral (6.6-7.5) to mildly alkaline (7.6-7.8).

Uses
The cornflower is considered a beneficial weed, and its edible flower can be used for culinary dec-
oration, for example to add colour to salads. In herbalism, a decoction of cornflower is effective in
treating conjunctivitis, and as a wash for tired eyes. Cornflowers have been used and prized histor-
ically for their blue pigment. Cornflowers are often used as an ingredient in some tea blends and
herbal teas, and is famous in the Lady Grey blend of Twinings.

Folklore and Symbolism

Logo of Conservative People’s Party of Estonia.

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78 Fundamentals of Weed Science

Flowering shoot of Cornflower. I. Disk-floret in vertical section.

In folklore, cornflowers were worn by young men in love; if the flower faded too quickly, it was
taken as a sign that the man’s love was not returned.

The blue cornflower has been the national flower of Estonia since 1968 and symbolizes daily
bread to Estonians. It is also the symbol of the Estonian political party, People’s Union, the Finn-
ish political party, National Coalition Party, and the Swedish political party, Liberal People’s
Party, and has since the dawn of the 20th century been a symbol for social liberalism there. It is
the official flower of the Swedish province of Östergötland and the school flower of Winchester
College and also of Dulwich College where it is said to have been the favourite flower of the
founder, Edward Alleyn.

The blue cornflower was one of the national symbols of Germany. This is partly due to the story
that when Queen Louise of Prussia was fleeing Berlin and pursued by Napoleon’s forces, she hid
her children in a field of cornflowers and kept them quiet by weaving wreaths for them from the
flowers. The flower thus became identified with Prussia, not least because it was the same color
as the Prussian military uniform. After the unification of Germany in 1871, it went on to become
a symbol of the country as a whole. For this reason, in Austria the blue cornflower is a political
symbol for pan-German and rightist ideas. Members of the Freedom Party wore it at the opening
of the Austrian parliament in 2006.

It was also the favourite flower of Louise’s son Kaiser Wilhelm I. Because of its ties to royalty, au-

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Types of Weed 79

thors such as Theodor Fontane have used it symbolically, often sarcastically, to comment on the
social and political climate of the time.

The cornflower is also often seen as an inspiration for the German Romantic symbol of the Blue
Flower.

Due to its traditional association with Germany, the cornflower has been made the official symbol
of the annual German-American Steuben Parade.

In France the Bleuet de France is the symbol of the 11th November 1918 armistice and, as such,
a common symbol for veterans (especially the now defunct poilus of World War I), similar to the
Remembrance poppies worn in the United Kingdom and in Canada.

The cornflower is also the symbol for motor neurone disease and amyotrophic lateral sclerosis.

Cornflowers are sometimes worn by Old Harrovians.

A Cornflower design (Blue Cornflower) was used by Corning Glass Works for the initial release of
Corning Ware Pyroceram Cookware. Its popularity in the United States, Canada, United Kingdom
and Australia was so high that it became the symbol of Corning Glass Works. The production of
Blue Cornflower Corning Ware (known as Pyrosil Ware & Pyroflam in Europe) spanned over 30
years, with initial production from 1958 to 1988 and a second short run from 1993-1994 marking
the 35th anniversary of their flagship consumer product. Corning Ware is no longer produced by
Corning Glass Works, as the Consumer Products Division was sold to Borden, Inc./World Kitch-
en LLC. in 1998. Production of Pyroceram was discontinued in the United States and Canada by
World Kitchen in 2000 in lieu of stoneware products, however, the popularity of the Blue Corn-
flower Corning Ware pattern (as well as a growing market for Pyroceram cookware) pushed World
Kitchen, LLC. into a re-introduce Pyroceram Cookware imported from France in 2009 with the
familiar Cornflower pattern.

In Paintings

Sergei Osipov. Cornflowers. 1976

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80 Fundamentals of Weed Science

Igor Grabar. Group portrait with cornflowers. 1914

Brassicaceae
Brassicaceae or Cruciferae is a medium-sized and economically important family of flowering
plants commonly known as the mustards, the crucifers, or the cabbage family.

The name Brassicaceae is derived from the included genus Brassica. The alternative older name,
Cruciferae, meaning “cross-bearing”, describes the four petals of mustard flowers, which resemble
a cross. Cruciferae is one of eight plant family names without the suffix -aceae that are authorized
alternative names (according to ICBN Art. 18.5 and 18.6 Vienna Code).

The family contains 372 genera and 4060 accepted species. The largest genera are Draba (440
species), Erysimum (261 species), Lepidium (234 species), Cardamine (233 species), and Alys-
sum (207 species).

The family contains the cruciferous vegetables, including species such as Brassica oleracea (e.g.,
broccoli, cabbage, cauliflower, kale, collards), Brassica rapa (turnip, Chinese cabbage, etc.), Bras-
sica napus (rapeseed, etc.), Raphanus sativus (common radish), Armoracia rusticana (horserad-
ish), Matthiola (stock) and the model organism Arabidopsis thaliana (thale cress).

Pieris rapae and other butterflies of the family Pieridae are some of the best-known pests of Bras-
sicaceae species planted as commercial crops.

Taxonomy
The family is included in the Brassicales according to the APG system. Older systems (e.g., Arthur
Cronquist’s) placed them into the Capparales, a now-defunct order that had a similar definition.

This family comprises about 365 genera and 3200 species all over the world; 94 species of 38 gen-
era are found in Nepal. The plants are mostly herbs. A close relationship has long been acknowl-
edged between the Brassicaceae and the caper family, Capparaceae, in part because members of
both groups produce glucosinolate (mustard oil) compounds. The Capparaceae as traditionally
circumscribed were paraphyletic with respect to Brassicaceae, with Cleome and several related
genera being more closely related to the Brassicaceae than to other Capparaceae. The APG II sys-
tem, therefore, has merged the two families under the name Brassicaceae. Other classifications

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Types of Weed 81

have continued to recognize the Capparaceae, but with a more restricted circumscription, either
including Cleome and its relatives in the Brassicaceae or recognizing them in the segregate family
Cleomaceae. The APG III system has recently adopted this last solution, but this may change as a
consensus arises on this point. This article deals with Brassicaceae sensu stricto, i.e. treating the
Cleomaceae and Capparaceae as segregated families.

Description

Aubrieta deltoidea (commonly known as purple rock cress) is a perennial wild flower used in gardening for its
ornamental large inflorescence.

The family consists mostly of herbaceous plants with annual, biennial, or perennial lifespans.
However, around the Mediterranean, they include also a dozen woody shrubs 1-3 m tall, e.g. in
northern Africa (Zilla spinosa and Ptilotrichum spinosum), in the Dalmatian islands (Dendralys-
sum and Cramboxylon), and chiefly in Canarias with some woody cruciferous genera: Dendros-
inapis, Descurainia, Parolinia, Stanleya, etc..

The leaves are alternate (rarely opposite), sometimes organized in basal rosettes; in rare shrubby
crucifers of Mediterranean their leaves are mostly in terminal rosettes, and may be coriaceous and
evergreen. They are very often pinnately incised and do not have stipules.

The structure of the flowers is extremely uniform throughout the family. They have four free sac-
cate sepals and four clawed free petals, staggered. They can be disymmetric or slightly zygomor-
phic, with a typical cross-like arrangement (hence the name Cruciferae). They have six stamens,
four of which are longer (as long as the petals) and are arranged in a cross like the petals and the
other two are shorter (tetradynamous flower). The pistil is made up of two fused carpels and the
style is very short, with two lobes. The ovary is superior. The flowers form ebracteate racemose
inflorescences, often apically corymb-like.

Pollination occurs by entomogamy; nectar is produced at the base of the stamens and stored on
the sepals.

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82 Fundamentals of Weed Science

Siliquae of Cardamine impatiens

The fruit is a peculiar kind of capsule named siliqua (plural siliquae). It opens by two valves, which
are the modified carpels, leaving the seeds attached to a framework made up of the placenta and
tissue from the junction between the valves (replum). Often, an indehiscent beak occurs at the top
of the style and one or more seeds may be borne there. Where a siliqua is less than three times as
long as it is broad, it is usually termed a silicula. The siliqua may break apart at constrictions oc-
curring between the segments of the seeds, thus forming a sort of loment (e.g., Raphanus), it may
eject the seeds explosively (e.g., Cardamine) or may be evolved in a sort of samara (e.g., Isatis).
The fruit is often the most important diagnostic character for plants in this family. Most members
share a suite of glucosinolate compounds that have a typical pungent odour usually associated with
cole crops.

Uses

Lunaria annua with ripe seed pods

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Types of Weed 83

Smelowskia americana is endemic to the midlatitude mountains of western North America.

The importance of this family for food crops has led to its selective breeding throughout history.
Some examples of cruciferous food plants are the cabbage, broccoli, cauliflower, turnip, rapeseed,
mustard, radish, horseradish, cress, wasabi, and watercress.

Matthiola (stock), Cheiranthus, Lobularia, and Iberis (candytufts) are appreciated for their flow-
ers. Lunaria (honesty) is cultivated for the decorative value of the translucent replum of the round
silicula that remains on the dried stems after dehiscence.

Capsella bursa-pastoris, Lepidium, and many Cardamine species are common weeds.

Isatis tinctoria (woad) was used in the past to produce the colour indigo.

Arabidopsis thaliana is a very important model organism in the study of the flowering plants (An-
giospermae).

Genera

• Aethionema • Lachnoloma
• Agallis • Leavenworthia
• Alliaria • Lepidium
• Alyssoides • Lepidostemon
• Alyssopsis • Leptaleum
• Alyssum • Lignariella
• Ammosperma • Lithodraba
• Anastatica • Lobularia
• Anchonium • Lonchophora

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84 Fundamentals of Weed Science

• Andrzeiowskia • Loxostemon
• Anelsonia • Lunaria
• Aphragmus • Lyocarpus
• Aplanodes • Lyrocarpa
• Arabidella • Macropodium
• Arabidopsis • Malcolmia
• Arabis • Mancoa
• Arcyosperma • Maresia
• Armoracia • Mathewsia
• Aschersoniodoxa • Matthiola
• Asperuginoides • Megacarpaea
• Asta • Megadenia
• Atelanthera • Menkea
• Athysanus • Menonvillea
• Aubrieta • Microlepidium
• Aurinia • Microsysymbrium
• Ballantinia • Microstigma
• Barbarea • Morettia
• Beringia • Moricandia
• Berteroa • Moriera
• Berteroella • Morisia
• Biscutella • Murbeckiella
• Bivonaea • Muricaria
• Blennodia • Myagrum
• Boechera • Nasturtiopsis
• Boleum • Nasturtium
• Boreava • Neomartinella
• Bornmuellera • Neotchihatchewia
• Borodinia • Neotorularia
• Botscantzevia • Nerisyrenia
• Brachycarpaea • Neslia
• Brassica • Nesocrambe
• Braya • Neuontobotrys
• Brayopsis • Notoceras
• Brossardia • Notothlaspi
• Bunias • Ochthodium
• Cakile • Octoceras
• Calepina • Olimarabidopsis

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Types of Weed 85

• Calymmatium • Onuris
• Camelina • Oreoloma
• Camelinopsis • Oreophyton
• Capsella • Ornithocarpa
• Cardamine • Orychophragmus
• Cardaminopsis • Otocarpus
• Cardaria • Oudneya
• Carinavalva • Pachycladon
• Carrichtera • Pachymitus
• Catadysia • Pachyphragma
• Catenulina • Pachypterygium
• Caulanthus • Parlatoria
• Caulostramina • Parodiodoxa
• Ceratocnemum • Parolinia
• Ceriosperma • Parrya
• Chalcanthus • Parryodes
• Chamira • Paysonia
• Chartoloma • Pegaeophyton
• Cheesemania • Peltaria
• Cheiranthus • Peltariopsis
• Chlorocrambe • Pennellia
• Chorispora • Petiniotia
• Christolea • Petrocallis
• Chrysobraya • Petrocallis
• Chrysochamela • Petroravenia
• Cithareloma • Phlebolobium
• Clastopus • Phlegmatospermum
• Clausia • Phoenicaulis
• Clypeola • Physaria
• Cochlearia • Physocardamum
• Coelonema • Physoptychis
• Coincya • Physorrhynchus
• Coluteocarpus • Platycraspedum
• Conringia • Polyctenium
• Cordylocarpus • Polypsecadium
• Coronopus • Pringlea
• Crambe • Prionotrichon
• Crambella • Pritzelago

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86 Fundamentals of Weed Science

• Cremolobus • Pseuderucaria
• Crucihimalaya • Pseudoarabidopsis
• Cryptospora • Pseudocamelina
• Cuphonotus • Pseudoclausia
• Cusickiella • Pseudofortuynia
• Cycloptychis • Pseudovesicaria
• Cymatocarpus • Psychine
• Cyphocardamum • Pterygiosperma
• Dactylocardamum • Pterygostemon
• Degenia • Pugionium
• Delpinophytum • Pycnoplinthopsis
• Descurainia • Pycnoplinthus
• Diceratella • Pyramidium
• Dichasianthus • Quezeliantha
• Dictyophragmus • Quidproquo
• Didesmus • Raffenaldia
• Didymophysa • Raphanorhyncha
• Dielsiocharis • Raphanus
• Dilophia • Rapistrum
• Dimorphocarpa • Reboudia
• Diplotaxis • Redowskia
• Dipoma • Rhammatophyllum
• Diptychocarpus • Rhizobotrya
• Dithyrea • Ricotia
• Dolichirhynchus • Robeschia
• Dontostemon • Rollinsia
• Douepea • Romanschulzia
• Draba • Roripella
• Drabastrum • Rorippa
• Drabopsis • Rytidocarpus
• Dryopetalon • Sameraria
• Eigia • Sarcodraba
• Elburzia • Savignya
• Enarthrocarpus • Scambopus
• Englerocharis • Schimpera
• Eremobium • Schivereckia
• Eremoblastus • Schizopetalon
• Eremodraba • Schlechteria

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Types of Weed 87

• Eremophyton • Schoenocrambe
• Ermania • Schouwia
• Ermaniopsis • Scoliaxon
• Erophila • Selenia
• Eruca • Sibara
• Erucaria • Sibaropsis
• Erucastrum • Silicularia
• Erysimum • Sinapidendron
• Euclidium • Sinapis
• Eudema • Sisymbrella
• Eutrema • Sisymbriopsis
• Euzomodendron • Sisymbrium
• Farsetia • Smelowskia
• Fezia • Sobolewskia
• Fibigia • Solms-laubachia
• Foleyola • Sophiopsis
• Fortuynia • Sphaerocardamum
• Galitzkya • Spirorhynchus
• Geococcus • Spryginia
• Glaribraya • Staintoniella
• Glastaria • Stanfordia
• Glaucocarpum • Stanleya
• Goldbachia • Stenopetalum
• Gorodkovia • Sterigmostemum
• Graellsia • Stevenia
• Grammosperma • Straussiella
• Guillenia • Streptanthella
• Guiraoa • Streptanthus
• Gynophorea • Streptoloma
• Halimolobos • Stroganowia
• Harmsiodoxa • Stubebdorffia
• Hedinia • Subularia
• Heldreichia • Succowia
• Heliophila • Synstemon
• Hemicrambe • Synthlipsis
• Hemilophia • Taphrospermum
• Hesperis • Tauscheria
• Heterodraba • Teesdalia

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88 Fundamentals of Weed Science

• Hirschfeldia • Teesdaliopsis
• Hollermayera • Tetracme
• Hormathophylla • Thellungiella
• Hornungia • Thelypodiopsis
• Hornwoodia • Thelypodium
• Hugueninia • Thlaspeocarpa
• Hymenolobus • Thlaspi
• Ianhedgea • Thysanocarpus
• Iberis • Trachystoma
• Idahoa • Trichotolinum
• Iodanthus • Trochiscus
• Ionopsidium • Tropidocarpum
• Irenepharsus • Turritis
• Isatis • Vella
• Ischnocarpus • Warea
• Iskandera • Weberbauera
• Iti • Werdermannia
• Ivania • Winklera
• Jundzillia • Xerodraba
• Kernera • Yinshania
• Kremeriella • Zerdana
• Lachnocapsa • Zilla

Cowpea
The cowpea (Vigna unguiculata) is one of several species of the widely cultivated genus Vigna.
Four subspecies are recognised, of which three are cultivated (more exist, including V. textilis, V.
pubescens, and V. sinensis):

Cowpeas are one of the most important food legume crops in the semiarid tropics covering Asia,
Africa, southern Europe, and Central and South America. A drought-tolerant and warm-weather
crop, cowpeas are well-adapted to the drier regions of the tropics, where other food legumes do
not perform well. It also has the useful ability to fix atmospheric nitrogen through its root nodules,
and it grows well in poor soils with more than 85% sand and with less than 0.2% organic matter
and low levels of phosphorus. In addition, it is shade tolerant, so is compatible as an intercrop
with maize, millet, sorghum, sugarcane, and cotton. This makes cowpeas an important component
of traditional intercropping systems, especially in the complex and elegant subsistence farming
systems of the dry savannas in sub-Saharan Africa. In these systems the haulm (dried stalks) of
cowpea is a valuable by-product, used as animal feed.

Research in Ghana found that selecting early generations of cowpea crops to increase yield is not
an effective strategy. Francis Padi from the Savannah Agricultural Research Institute in Tamale,

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Types of Weed 89

Ghana, writing in Crop Science, suggests other methods such as bulk breeding are more efficient
in developing high-yield varieties.

Cowpea flower

According to the USDA food database, the leaves of the cowpea plant have the highest percentage
of calories from protein among vegetarian foods.

Taxonomy and Etymology


Vigna unguiculata is a member of the Vigna (peas or beans) genus. Unguiculata is Latin for “with
a small claw”, which reflects the small stalks on the flower petals. All cultivated cowpeas are found
within the universally accepted V. unguiculata subspecies unguiculata classification, which is
then commonly divided into four cultivar groups: Unguiculata, Biflora, Sesquipedalis, and Tex-
tilis. Some well-known common names for cultivated cowpeas include Lesera/ Dangbodi (লেছেৰা/
ডাংবডি) in Assamese, Behlawi in Mizo, black-eye pea, southern pea, yardlong bean, catjang and
Crowder Pea. The classification of the wild relatives within V. unguiculata is more complicated,
with over 20 different names having been used and between 3 and 10 subgroups described. The
original subgroups of stenophylla, dekindtiana and tenuis appear to be common in all taxonomic
treatments, with the earlier described variations pubescens and protractor being raised to sub
species level by a 1993 charactisation.

Common names of Vigna unguiculata unguiculata cultivar groups

Group Common name

Unguiculata crowder-pea, southern pea, black-eyed pea

Biflora catjang, sow-pea

Sesquipedalis yardlong bean, asparagus bean, Chinese long-bean

Textilis

The first written reference using cowpea appeared in 1798 in the United States. The name was
most likely acquired due to their use as a fodder crop for cows. The common name of black-eyed
pea, used for the unguiculata cultivar group, describes the presence of a distinctive black spot at

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90 Fundamentals of Weed Science

the hilum of the seed. Black-eyed peas were first introduced to the southern states in the United
States and some early varieties had peas squashed closely together in their pods, leading to the
other common names of southern pea and crowder-pea. Sesquipedalis in Latin means “foot and
a half long”, and this subspecies which arrived in the United States via Asia is characterised by
unusually long pods, leading to the common names of yardlong bean, asparagus bean and Chinese
long-bean. In West Africa the plant is named niebe, wake or ewa.

Description
There is a large morphological diversity found within the crop, and the growth conditions and
grower preferences for each variety vary from region to region.

History
Although there is no archaeological evidence for early cowpea cultivation the centre of diversity of
the cultivated cowpea is West Africa, leading to the current consensus that this is the likely cen-
tre of origin and place of early domestication. While it may be uncertain when cultivation began,
remains of charred cowpeas from rock shelters in Central Ghana have been dated to the second
millennium BCE. In 2300 BC the cowpea is believed to have made its way into South East Asia
where secondary domestication events may have occurred. The first written references to the cow-
pea were in 300BC and they probably reached Central and North America during the slave trade
through the 17thto early 19th centuries.

Cultivation
Cowpeas are grown mostly for their edible beans, although the leaves, green peas and green pea
pods can also be consumed, meaning the cowpea can be used as a food source before the dried peas
are harvested. Cowpeas thrive in poor dry conditions, growing well in soils up to 85% sand. This
makes them a particularly important crop in arid, semi-desert regions where not many other crops
will grow. As well as an important source of food for humans in poor arid regions the crop can also
be used as feed for livestock. This predominately occurs in India, where the stock is fed cowpea as
forage or fodder. The nitrogen fixing ability means that as well as functioning as a sole-crop, the
cowpea can be effectively intercropped with sorghum, millet, maize, cassava or cotton.

Pests and Diseases


Insects are a major factor in the low yields of African cowpea crops, and they affect each tissue
component and developmental stage of the plant. In bad infestations insect pressure is responsible
for over 90% loss in yield. The legume pod borer Maruca (testulalis) vitrata, is the main pre-har-
vest pest of the cowpea. It causes damage to the flower buds, flowers and pods of the plant. Oth-
er important pests include pod sucking bugs, thrips and the post-harvest weevil Callosobruchus
maculatus.

Culinary use
In Tamil Nadu, India, between the Tamil months of Maasi (February) and Panguni (March), a
cake-like dish called kozhukattai (steamed sweet dumplings – also called Sukhiyan in Kerala) is

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Types of Weed 91

prepared with cooked and mashed cowpeas mixed with jaggery, ghee, and other ingredients. That-
ta payir in sambar and pulikkuzhambu (spicy semisolid gravy in tamarind paste) is a popular dish
in Tamil Nadu.

In Sri Lanka, cowpeas (කවුපි in Sinhala) are cooked in many different ways, one of which is with
coconut milk.

In Turkey, cowpeas can be lightly boiled, covered with olive oil, salt, thyme, and garlic sauce, and
eaten as an appetizer; they are cooked with garlic and tomatoes; and they can be eaten in bean salad.

NCT Dream, a popular Korean pop band, used cowpeas in making ricecakes for Chuseok, a Korean
holiday.

Nutrition and Health


Cowpeas provide a rich source of proteins and calories, as well as minerals and vitamins. A cowpea
seed can consist of 25% protein and is low in anti-nutritional factors. This diet complements the
mainly cereal diet in countries that grow cowpeas as a major food crop.

Production and Consumption


Most cowpeas are grown on the African continent, particularly in Nigeria and Niger which account
for 66% of world cowpea production. The Sahel region also contains other major producers such
as Burkina Faso, Ghana, Senegal and Mali. Niger is the main exporter of cowpeas and Nigeria the
main importer. Exact figures for cowpea production are hard to come up with as it is not a major
export crop. A 1997 estimate suggests that cowpeas are cultivated on 12.5 million hectares and
have a worldwide production of 3 million tonnes. While they play a key role in subsistence farming
and livestock fodder, the cowpea is also seen as a major cash crop by Central and West African
farmers, with an estimated 200 million people consuming cowpea on a daily basis.

According to the Food and Agriculture Organisation of the United Nations (FAO), as of 2012, the
average cowpea yield in Western Africa was an estimated 483 kg/ha, which is still 50% below the
estimated potential production yield. In some tradition cropping methods the yield can be as low
as 100 kg/ha.

Outside Africa, the major production areas are Asia, Central America, and South America. Brazil
is the world’s second-leading producer of cowpea seed, producing 600,000 tonnes annually. The
amount of protein content of cowpea’s leafy parts consumed annually in Africa and Asia is equiva-
lent to 5 million tonnes of dry cowpea seeds, representing as much as 30% of the total food legume
production in the lowland tropics.

Solanaceae
The Solanaceae, or nightshades, are an economically important family of flowering plants. The
family ranges from annual and perennial herbs to vines, lianas, epiphytes, shrubs, and trees, and
includes a number of important agricultural crops, medicinal plants, spices, weeds, and ornamen-
tals. Many members of the family contain potent alkaloids, and some are highly toxic, but many
cultures eat nightshades, in some cases as staple foods. The family belongs to the order Solanales,

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92 Fundamentals of Weed Science

in the asterid group and class Magnoliopsida (dicotyledons). The Solanaceae consists of about 98
genera and some 2,700 species, with a great diversity of habitats, morphology and ecology.

Fruits including tomatoes, tomatillos, eggplant, and chili peppers.

The name Solanaceae derives from the genus Solanum, “the nightshade plant”. The name may
come from a perceived resemblance of certain solanaceous flowers to the sun and its rays. At least
one species of Solanum is known as the “sunberry”. Alterna-tively, the name could originate from
the Latin verb solari, meaning “to soothe”, presumably refer-ring to the soothing pharmacological
properties of some of the psychoactive species of the family.

The family has a worldwide distribution, being present on all continents except Antarctica. The
greatest diversity in species is found in South America and Central America.

The Solanaceae include a number of commonly collected or cultivated species. The most econom-
ically important genus of the family is Solanum, which contains the potato (S. tuberosum, in fact,
another common name of the family is the “potato family”), the tomato (S. lycopersicum), and the
eggplant or aubergine (S. melongena). Another important genus, Capsicum, produces both chili
peppers and bell peppers.

The genus Physalis produces the so-called groundcherries, as well as the tomatillo (Physalis phila-
delphica), the Cape gooseberry and the Chinese lantern. The genus Lycium contains the boxthorns
and the wolfberry Lycium barbarum. Nicotiana contains, among other species, tobacco. Some
other important members of Solanaceae include a number of ornamental plants such as Petunia,
Browallia, and Lycianthes, and sources of psychoactive alkaloids, Datura, Mandragora (man-
drake), and Atropa belladonna (deadly nightshade). Certain species are widely known for their
medicinal uses, their psychotropic effects, or for being poisonous.

Most of the economically important genera are contained in the subfamily Solanoideae, with the
exceptions of tobacco (Nicotiana tabacum, Nicotianoideae) and petunia (Petunia × hybrida, Pe-
tunioideae),

Many of the Solanaceae, such as tobacco and petunia, are used as model organisms in the investi-
gation of fundamental biological questions at the cellular, molecular, and genetic levels.

Description
Plants in the “Solanaceae” can take the form of herbs, shrubs, trees, vines and lianas, and some-

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Types of Weed 93

times epiphytes. They can be annuals, biennials, or perennials, upright or decumbent. Some have
subterranean tubers. They do not have laticifers, nor latex, nor coloured saps. They can have a
basal or terminal group of leaves or neither of these types. The leaves are generally alternate or
alternate to opposed (that is, alternate at the base of the plant and opposed towards the inflores-
cence). The leaves can be herbaceous, leathery, or transformed into spines. The leaves are general-
ly petiolate or subsessile, rarely sessile. They are frequently inodorous, but on occasions, they are
aromatic or fetid. The foliar lamina can be either simple or compound, and the latter can be either
pinnatifid or ternate. The leaves have reticulated venation and lack a basal meristem. The laminae
are generally dorsiventral and lack secretory cavities. The stomata are generally confined to one of
a leaf’s two sides; they are rarely found on both sides.

Illustration of Solanum dulcamara, 1.- Flower, 2.- Flower in longitudinal section, without the petals; 3.- Androecium;
4.- Ovary, in transverse section; 5.- Seed viewed from above; 6.- Seed in transverse section, note the curved embryo
surrounding the endosperm; A.- Branch with leaves and flowers; B.- Stem with immature and mature fruit

The flowers are generally hermaphrodites, although some are monoecious, andromonoecious, or
dioecious species (such as some Solanum or Symonanthus). Pollination is entomophilous. The
flowers can be solitary or grouped into terminal, cymose, or axillary inflorescences. The flowers
are medium-sized, fragrant (Nicotiana), fetid (Anthocercis), or inodorous. The flowers are usually
actinomorphic, slightly zygomorphic, or markedly zygomorphic (for example, in flowers with a
bilabial corolla in Schizanthus species). The irregularities in symmetry can be due to the androe-
cium, to the perianth, or both at the same time. In the great majority of species, the flowers have
a differentiated perianth with a calyx and corolla (with five sepals and five petals, respectively) an
androecium with five stamens and two carpels forming a gynoecium with a superior ovary (they
are therefore referred to as pentamers and tetracyclic). The stamens are epipetalous and are typi-
cally present in multiples of four or five, most commonly four or eight. They usually have a hypog-
ynous disk. The calyx is gamosepalous (as the sepals are joined together forming a tube), with the
(4)5(6) segments equal, it has five lobes, with the lobes shorter than the tube, it is persistent and

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94 Fundamentals of Weed Science

often accrescent. The corolla usually has five petals that are also joined together forming a tube.
Flower shapes are typically rotate (wheel-shaped, spreading in one plane, with a short tube) or
tubular (elongated cylindrical tube), campanulated or funnel-shaped.

The androecium has (2)(4)5(6) free stamens within it, oppositsepals (that is, they alternate with the
petals), they are usually fertile or, in some cases (for example in Salpiglossideae) they have stami-
nodes. In the latter case, there is usually either one staminode (Salpiglossis) or three (Schizanthus).
The anthers touch on their upper end forming a ring, or they are completely free, dorsifixed, or ba-
sifixed with poricide dehiscence or through small longitudinal cracks. The stamen’s filament can be
filliform or flat. The stamens can be inserted inside the coralline tube or exserted. The plants demon-
strate simultaneous microsporogenesis, the microspores are tetrad, tetrahedral, or isobilateral. The
pollen grains are bicellular at the moment of dehiscence, usually open and angular.

The gynoecium is bicarpelar (rarely three- or five-locular) with a superior ovary and two locules,
which may be secondarily divided by false septa, as is the case for Nicandreae and Datureae. The
gynoecium is located in an oblique position relative to the flower’s median plane. They have one
style and one stigma; the latter is simple or bilobate. Each locule has one to 50 ovules that are ana-
tropous or hemianatropous with axillar placentation. The development of the embryo sack can be
the same as for Polygonum or Allium species. The embryo sack’s nuclear poles become fused be-
fore fertilization. The three antipodes are usually ephemeral or persistent as in the case of Atropa.
The fruit can be a berry as in the case of the tomato or wolfberry a dehiscent capsule as in Datura,
or a drupe. The fruit has axial placentation. The capsules are normally septicidal or rarely locu-
licidal or valvate. The seeds are usually endospermic, oily (rarely starchy), and without obvious
hairs. The seeds of most Solanaceae are round and flat, about 2–4 mm (0.079–0.157 in) in diame-
ter. The embryo can be straight or curved, and has two cotyledons. Most species in the Solanaceae
have 2n=24 chromosomes, but the number may be a higher multiple of 12 due to polyploidy. Wild
potatoes, of which there are about 200, are predominantly diploid (2 × 12 = 24 chromosomes),
but triploid (3 × 12 = 36 chromosomes), tetraploid (4 × 12 = 48 chromosomes), pentaploid (5 × 12
= 60) and even hexaploid (6 × 12 = 72 chromosome) species or populations exist. The cultivated
species Solanum tuberosum has 4 × 12 = 48 chromosomes. Some Capsicum species have 2 × 12 =
24 chromosomes, while others have 26 chromosomes.

Diversity of Characteristics
Despite the previous description, the Solanaceae exhibit a large morphological variability, even in
their reproductive characteristics. Examples of this diversity include:

• The number of carpels that form the gynoecium

In general, the Solanaceae have a gynoecium (the female part of the flower) formed of two car-
pels. However, Melananthus has a monocarpelar gynoecium, there are three or four carpels in
Capsicum, three to five in Nicandra, some species of Jaborosa and Trianaea and four carpels in
Iochroma umbellatum.

• The number of locules in the ovary

The number of locules in the ovary is usually the same as the number of carpels. However, some

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Types of Weed 95

species occur in which the numbers are not the same due to the existence of false septa (internal
walls that subdivide each locule), such as in Datura and some members of the Lycieae (the genera
Grabowskia and Vassobia).

• Type of ovules and their number

The ovules are generally inverted, folded sharply backwards (anatropous), but some genera have
ovules that are rotated at right angles to their stalk (campilotropous) as in Phrodus, Grabows-
kia or Vassobia), or are partially inverted (hemitropous as in Cestrum, Capsicum, Schizanthus
and Lycium). The number of ovules per locule also varies from a few (two pairs in each locule in
Grabowskia, one pair in each locule in Lycium) and very occasionally only one ovule is in each
locule as for example in Melananthus.

• The type of fruit

The fruits of the great majority of the Solanaceae are berries or capsules (including pyxidia) and less
often drupes. Berries are common in the subfamilies Cestroideae, Solanoideae (with the exception of
Datura, Oryctus, Grabowskia and the tribe Hyoscyameae) and the tribe Juanulloideae (with the ex-
ception of Markea). Capsules are characteristic of the subfamilies Cestroideae (with the exception of
Cestrum) and Schizanthoideae, the tribes Salpiglossoideae and Anthocercidoideae, and the genus Da-
tura. The tribe Hyoscyameae has pyxidia. Drupes are typical of the Lycieae tribe and in Iochrominae.

Alkaloids
Alkaloids are nitrogenous organic substances produced by plants as a secondary metabolite and
which have an intense physiological action on animals even at low doses. Solanaceae are known for
having a diverse range of alkaloids. To humans, these alkaloids can be desirable, toxic, or both. The
tropanes are the most well-known of the alkaloids found in the Solanaceae. The plants that con-
tain these substances have been used for centuries as poisons. However, despite being recognized
as poisons, many of these substances have invaluable pharmaceutical properties. Many species
contain a variety of alkaloids that can be more or less active or poisonous, such as scopolamine, at-
ropine, hyoscyamine, and nicotine. They are found in plants such as the henbane (Hyoscyamus al-
bus), belladonna (Atropa belladonna), datura or jimson (Datura stramonium), mandrake (Man-
dragora autumnalis), tobacco, and others. Some of the main types of alkaloids are:

Chemical structure of solanine

• Solanine: A toxic glycoalkaloid with a bitter taste, it has the formula C45H73NO15. It is formed
by the alkaloid solanidine with a carbohydrate side chain. It is found in leaves, fruit, and
tubers of various Solanaceae such as the potato and tomato. Its production is thought to
be an adaptive defence strategy against herbivores. Substance intoxication from solanine
is characterized by gastrointestinal disorders (diarrhoea, vomiting, abdominal pain) and

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96 Fundamentals of Weed Science

neurological disorders (hallucinations and headache). The median lethal dose is between 2
and 5 mg per kg of body weight. Symptoms become manifest 8 to 12 hr after ingestion. The
amount of these glycoalkaloids in potatoes, for example, varies significantly depending of
environmental conditions during their cultivation, the length of storage, and the variety.
The average glycoalkaloid concentration is 0.075 mg/g of potato. Solanine has occasionally
been responsible for poisonings in people who ate berries from species such as Solanum
nigrum or Solanum dulcamara, or green potatoes.

Chemical structure of the tropanes.

• Tropanes: The term “tropane” comes from a genus in which they are found, Atropa (the bella-
donna genus). Atropa is named after the Greek Fate, Atropos, who cut the thread of life. This
nomenclature reflects its toxicity and lethality. They are bicyclic organic nitrogen compounds
(IUPAC nomenclature: 8-Methyl-8-azabicyclo[3.2.1]octane), with the chemical formula of
C8H15N. These alkaloids include, among others, atropine, cocaine, scopolamine, and hyoscy-
amine. They are found in various species, such as mandrake (Mandragora autumnalis), black
henbane or stinking nightshade (Hyoscyamus niger), belladonna (Atropa belladonna) the
stramonium (Datura stramonium) and Brugmansia species, as well as many others in the
Solanaceae family. Pharmacologically, they are the most powerful known anticholinergics in
existence, meaning they inhibit the neurological signals transmitted by the endogenous neu-
rotransmitter, acetylcholine. More commonly, they can halt many types of allergic reactions.
Symptoms of overdose may include dry mouth, dilated pupils, ataxia, urinary retention, hallu-
cinations, convulsions, coma, and death. Atropine, a commonly used ophthalmological agent,
dilates the pupils and thus facilitates examination of the interior of the eye. In fact, juice from
the berries of A. belladonna were used by Italian courtesans during the Renaissance to exag-
gerate the size of their eyes by causing the dilation of their pupils. Despite the extreme toxicity
of the tropanes, they are useful drugs when administered in extremely small dosages. They
can reverse cholinergic poisoning, which can be caused by overexposure to organophosphate
insecticides and chemical warfare agents such as sarin and VX. Scopolamine (found in Hyos-
cyamus muticus and Scopolia atropioides), is used as an antiemetic against motion sickness
or for people suffering from nausea as a result of receiving chemotherapy. Scopolamine and
hyoscyamine are the most widely used tropane alkaloids in pharmacology and medicine due
to their effects on the parasympathetic nervous system. Atropine has a stimulant effect on the
central nervous system and heart, whereas scopolamine has a sedative effect. These alkaloids

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Types of Weed 97

cannot be substituted by any other class of compounds, so they are still in demand. This is one
of the reasons for the development of an active field of research into the metabolism of the
alkaloids, the enzymes involved, and the genes that produce them. Hyoscyamine 6-β hydrox-
ylase, for example, catalyses the hydroxylation of hyoscyamine that leads to the production of
scopolamine at the end of the tropane’s biosynthetic pathway. This enzyme has been isolated
and the corresponding gene cloned from three species: H. niger, A. belladonna and B. candida.

Chemical structure of nicotine.

• Nicotine: Nicotine (IUPAC nomenclature (S)-3-(1-methylpyrrolidin-2-il) pyridine) is


a pyrrolidine alkaloid produced in large quantities in the tobacco plant (Nicotiana ta-
bacum), but is also found in lower concentrations in other species such as the potato,
tomato, and pepper. Its function in a plant is to act as a defence against herbivores, as
it is an excellent neurotoxin, in particular against insects. In fact, nicotine has been
used for many years as an insecticide, although its use is currently being replaced by
synthetic molecules derived from its structure. At low concentrations, nicotine acts as
a stimulant in mammals, which causes the dependency in smokers. Like the tropanes,
it acts on cholinergic neurons, but with the opposite effect (it is an agonist as opposed
to an antagonist). It has a higher specificity for nicotinic acetylcholine receptors than
other ACh proteins.

Chemical structure of capsaicin

• Capsaicin: Capsaicin (IUPAC nomenclature 8-methyl-N-vanillyl-trans-6-nonenamide) is


structurally different from nicotine and the tropanes. It is found in species of the genus
Capsicum, which includes chilies and habaneros and it is the active ingredient that de-
termines the Scoville rating of these spices. The compound is not noticeably toxic to hu-
mans. However, it stimulates specific pain receptors in the majority of mammals, specifi-
cally those related to the perception of heat in the oral mucosa and other epithelial tissues.
When capsaicin comes into contact with these mucosae, it causes a burning sensation little
different from a burn caused by fire. Capsaicin affects only mammals, not birds. Pepper
seeds can survive the digestive tracts of birds; their fruit becomes brightly coloured once its

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98 Fundamentals of Weed Science

seeds are mature enough to germinate, thereby attracting the attention of birds that then
distribute the seeds. Capsaicin extract is used to make pepper spray, a useful deterrent
against aggressive mammals.

Distribution

Map showing the distribution of the Solanaceae throughout the world (light green areas)

Even though members of the Solanaceae are found on all continents except Antarctica, the greatest
variety of species are found in Central America and South America. Centers of diversity also occur
in Australia and Africa. Solanaceae occupy a great number of different ecosystems, from deserts
to rainforests, and are often found in the secondary vegetation that colonizes disturbed areas. In
general, plants in this family are of tropical and temperate distribution.

Taxonomy
The following taxonomic synopsis of the solanaceas, including subfamilies, tribes and genera, is
based on the most recent molecular phylogenetics studies of the family:

Cladogram showing the relationship between the three genera of the Solanaceae family

Cestroideae (Browallioideae)

Cestrum elegans, a cestroidea used as an ornamental

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Types of Weed 99

Browallia americana

Flower of Salpiglossis sinuata, Botanischer Garten Jena, Germany

This subfamily is characterised by the presence of pericyclic fibres, an androecium with four or five
stamens, frequently didynamous. The basic chromosome numbers are highly variable, from x=7
to x=13. The subfamily consists of eight genera (divided into three tribes) and about 195 species
distributed throughout the Americas. The Cestrum genus is the most important, as it contains 175
of the 195 species in the subfamily. The Cestreae tribe is unusual because it includes taxa with long
chromosomes (from 7.21 to 11.511 µm in length), when the rest of the family generally possesses
short chromosomes (for example between 1.5 and 3.52 µm in the Nicotianoideae)
• Browallieae Hunz.

o Browallia L., genus with six species distributed throughout the neotropic ecozone
to Arizona in the United States

o Streptosolen Miers, monotypic genus native to the Andes

• Cestreae tribe Don, three genera of woody plants, generally shrubs

o Cestrum L., some 175 species distributed throughout the neotropic ecozone

o Sessea Ruiz & Pav., 16 species from the Andes

o Vestia Willd., monotypic genus from Chile

• Salpiglossideae tribe (Benth.) Hunz.

o Reyesia Gay, four species, distributed throughout Argentina and Chile

o Salpiglossis Ruiz & Pav., two species originating from southern South America

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100 Fundamentals of Weed Science

Goetzeoideae
This subfamily is characterized by the presence of drupes as fruit and seeds with curved embryos
and large fleshy cotyledons. The basic chromosome number is x=13. It includes four genera and
five species distributed throughout the Greater Antilles. Some authors suggest their molecular
data indicate the monotypic genera Tsoala Bosser & D’Arcy should be included in this subfamily,
endemic to Madagascar, and Metternichia to the southeast of Brazil. Goetzeaceae Airy Shaw is
considered as a synonym of this subfamily.
• Coeloneurum Radlk., monotypic genus endemic to Hispaniola
• Espadaea Rchb., monotypic, from Cuba
• Goetzea Wydler, includes two species from the Antilles
• Henoonia Griseb., monotypic, originating in Cuba

Petunioideae
Molecular phylogenetics indicates that Petunioideae is the sister clade of the subfamilies with chro-
mosome number x=12 (Solanoideae and Nicotianoideae). They contain calistegins, alkaloids similar
to the tropanes. The androecium is formed of four stamens (rarely five), usually with two different
lengths. The basic chromosome number of this subfamily can be x=7, 8, 9 or 11. It consists of 13 gen-
era and some 160 species distributed throughout Central and South America. Molecular data suggest
the genera originated in Patagonia. Benthamiella, Combera, and Pantacantha form a clade that can
be categorized as a tribe (Benthamielleae) that should be in the subfamily Goetzeoideae.

Nierenbergia frutescens, a petunoidea.

• Benthamiella Speg., 12 species native to Patagonia

• Bouchetia Dunal, three neotropical species

• Brunfelsia L., around 45 species from the neotropics

• Combera Sandw., two species from Patagonia

• Fabiana Ruiz & Pav., 15 species native to the Andes

• Hunzikeria D’Arcy, three species from the southwest United States and Mexico

• LatuaPhil., one species from the south of Chile

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Types of Weed 101

• Leptoglossis Benth., seven species from western South America

• Nierembergia Ruiz & Pav., 21 species from South America

• Pantacantha Speg., monospecific genus from Patagonia

• Calibrachoa Cerv. ex La Llave & Lex. consists of 32 species from the neotropics. The mor-
phological data suggest this genus should be included within the Petunia. However, the
molecular and cytogenetic data indicate both should be kept separate. In fact, Calibrachoa
has a basic chromosome number x=9, while that of Petunia is x=7.

• Petunia (Juss.) Wijsman, 18 species from South America

• Plowmania Hunz. & Subils, monotypic genus from Mexico and Guatemala

Schizanthoideae

Zygomorphic flowers, with bilabiate corolla of Schizanthus pinnatus, a schizanthoidea ornamental

The Schizanthoideae include annual and biennial plants with tropane alkaloids, without pericyclic
fibres, with characteristic hair and pollen grains. The flowers are zygomorphic. The androecium
has two stamens and three stamenodes, anther dehiscence is explosive. The embryo is curved. The
basic chromosome number is x=10. Schizanthus is a somewhat atypical genus among the Solana-
ceae due to its strongly zygomorphic flowers and basic chromosome number. Morphological and
molecular data suggest Schizanthus is a sister genus to the other Solanaceae and diverged early
from the rest, probably in the late Cretaceous or in the early Cenozoic, 50 million years ago. The
great diversity of flower types within Schizanthus has been the product of the species’ adaptation
to the different types of pollinators that existed in the Mediterranean, high alpine, and desert eco-
systems then present in Chile and adjacent areas of Argentina.

• Schizanthus Ruiz & Pav., 12 species originating from Chile.

Schwenckioideae
Annual plants with pericyclic fibres, their flowers are zygomorphic, the androecium has four didy-

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102 Fundamentals of Weed Science

namous stamens or three stamenodes; the embryo is straight and short. The basic chromosome
number is x=12. It includes four genera and some 30 species distributed throughout South Amer-
ica.

• Heteranthia Nees & Mart., one species from Brazil

• Melananthus Walp., five species from Brazil, Cuba, and Guatemala

• Protoschwenckia Soler , monotypic genus from Bolivia and Brazil, some molecular phylo-
genetic studies have suggested this genus has an uncertain taxonomic position within the
subfamily

• Schwenckia L., 22 species distributed throughout the neotropical regions of America

Nicotianoideae
• Anthocercideae G. Don: This tribe, endemic to Australia, contains 31 species in seven gen-
era. Molecular phylogenetic studies of the tribe indicate it is the sister of Nicotiana, and
the genera Anthocercis, Anthotroche, Grammosolen, and Symonanthus are monophyletic.
Some characteristics are also thought to be derived from within the tribe, such as the uni-
locular stamens with semicircular opercula, bracteolate flowers, and berries as fruit.

Tobacco inflorescence, Nicotiana tabacum

o Anthocercis Labill., 10 species, Australia

o Anthotroche Endl., four species, Australia

o Crenidium Haegi, monotypic genus, Australia

o Cyphanthera Miers, 9 species, Australia

o Duboisia R.Br., four species, Australia

o Gramnosolen Haegi, two species, Australia

o Symonanthus Haegi, two species, Australia

• Nicotianeae tribe Dum.

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Types of Weed 103

o Nicotiana L., genus widely distributed, with 52 American species, 23 Australian,


and one African

Solanoideae

Capsicum frutescens cultivar “tabasco”, a solanoidea

Flor de beleño (Hyoscyamus niger)

Iochroma australe flower

• Capsiceae Dumort

o Capsicum L. includes 40 accepted neotropical species

o Lycianthes (Dunal) Hassler, some 200 species distributed throughout America and
Asia

• Datureae G. Don, two genera are perfectly differentiated at both the morphological and mo-
lecular levels, Brugmansia includes tree species, while Datura contains herbs or shrubs,
the latter genus can be divided into three sections: Stramonium, Dutra and Ceratocaulis.

o Brugmansia Persoon, six species from the Andes

o Datura L., 12 neotropical species

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104 Fundamentals of Weed Science

• Hyoscyameae Endl.

o Anisodus Link, four species from China, India and the Himalayas

o Atropa L., four Euro-Asiatic species

o Atropanthe Pascher, monotypic genus from China

o Hyoscyamus L., 10 accepted species distributed from the Mediterranean to China

o Physochlaina G. Don, 6 accepted Euro-Asiatic species

o Przewalskia Maxim., one species from China

o Scopolia Jacq., disjointed distribution with one European species and another from
Japan

• Jaboroseae Miers

o Jaborosa Juss., genus that includes 23 species from South America.

• Solandreae Miers

o Subtribe Juanulloinae consists 10 genera of trees and epiphytic shrubs with a neo-
tropical distribution . Some of these genera (Dyssochroma, Merinthopodium and
Trianaea) show a clear dependency on various species of bats both for pollination
and dispersion of seeds.

 Dyssochroma Miers, two species from the south of Brazil

 Ectozoma Miers

 Hawkesiophyton Hunz.

 Juanulloa Ruiz & Pav., 11 species from South and Central America

 Markea Rich., 9 species from South and Central America

 Merinthopodium J. Donn. Sm. three species originating from South Amer-


ica

 Rahowardiana D’ Arcy

 Schultesianthus Hunz., eight neotropical species

 Trianaea Planch. & Linden, six South American species

o Subtribe Solandrinae, a monotypical subtribe, differs from Juanulloinae in that its


embryos have incumbent cotyledons and semi-inferior ovaries.

o Solandra Sw., 10 species from the neotropical regions of America

• Lycieae Hunz. has three genera of woody plants, which grow in arid or semiarid climates.

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Types of Weed 105

The cosmopolitan genus Lycium is the oldest in the tribe and it has the greatest morpho-
logical variability. Molecular phylogenetic studies suggest both Grabowskia and Phrodus
should be included in the Lycium, and this genus, along with Nolana and Sclerophylax,
form a clade (Lyciina), which currently lacks a taxonomic category. The red fleshy berries
dispersed by birds are the main type of fruit in Lycium. The different types of fruit in this
genus have evolved from the type of berry just mentioned to a drupe with a reduced num-
ber of seeds.

o Grabowskia Schltdl., three species from South America

o Lycium L., 83 cosmopolitan species

o Phrodus Miers, two species endemic to the north of Chile

• Mandragoreae (Wettst.) Hunz. & Barboza tribe does not have a defined systematic position
according to molecular phylogenetic studies.

o Mandragora L., two species from Eurasia

• Nicandreae Wettst. is a tribe with two South American genera. Molecular phylogenetic
studies indicate the genera are not interrelated nor are they related with other genera of the
family, so their taxonomic position is uncertain.

o Exodeconus Raf., six species from western South America

o Nicandra Adans, one species distributed throughout neotropical regions

• Nolaneae Rchb. are mostly herbs and small shrubs with succulent leaves, they have very
beautiful flowers that range from white to various shades of blue, their fruit is schizocarpal,
giving rise to various nuts.

o Nolana L., 89 species distributed throughout western South America

• Physaleae Miers, is a large tribe that is the sister of Capsiceae.

o Subtribe Iochrominae (Miers) Hunz., a clade within the Physaleae tribe. contains
37 species, mainly distributed in the Andes, assigned to six genera. The members of
this subtribe are characterized by being woody shrubs or small trees with attractive
tubular or rotated flowers. They also possess great floral diversity, containing every
type is present in the family. Their flowers can be red, orange, yellow, green, blue,
purple, or white. The corolla can be tubular to rotated, with a variation of up to
eight times in the length of the tube between the various species.

 Acnistus Schott, one species distributed throughout the neotropics

 Dunalia Kunth., five species from the Andes

 Iochroma Benth., 24 species from the Andes

 Saracha Ruiz & Pav., two species from the Andes.

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106 Fundamentals of Weed Science

 Vassobia Rusby, two South American species

 Eriolarynx Hunz., three species from Argentina and Bolivia

o Physalinae (Miers) Hunz. , a monophyletic subtribe, contains 10 genera and in-


cludes herbs or woody shrubs with yellow, white, or purple solitary axillary flowers
pollinated by bees. Once pollination occurs, the corolla falls and the calyx expands
until it entirely covers the boll that is developing (the calyx is called accrescent).
In many species, the calyx turns yellow or orange on maturity. The berries contain
many greenish to yellow-orange seeds, often with red or purple highlights.

 Brachistus Miers, three species from Mexico and Central America

 Chamaesaracha (A.Gray) Benth. & Hook., has 10 species from Mexico and
Central America.

 Leucophysalis Rydberg, includes 3 species from the south west of the Unit-
ed States and Mexico.

 Margaranthus Schlecht., with 1 species from Mexico.

 Oryctes S. Watson, monotypic genus from the south west of the United States.

 Quincula Raf. with just 1 species from the south west of the United States
and from Mexico.

 Physalis L., the largest genus of the subtribe, with 85 species distributed
through the tropical regions of the Americas and with 1 species in China.

 Witheringia L’ Heritier, genus with 15 species from neotropical regions.

 Tzeltalia, genus segregated from Physalis, with 2 species distributed


throughout Mexico and Guatemala.

 Darcyanthus, genus with just 1 species originating in Bolivia and Peru.

o Subtribe Salpichroinae, this is a subtribe of Physaleae that includes 16 American


species distributed in 1 genera:

 Nectouxia Kunth., monotypic genus that is endemic to Mexico.

 Salpichroa Miers, genus with 15 species from the Andes and other regions
of South America.

o Subtribe Withaninae, is a subtribe of Physaleae with a broad distribution, including


9 genera:

 Archiphysalis Kuang, with 3 species from China and Japan.

 Athenaea Sendtn., which includes 7 species from Brazil.

 Aureliana Sendtn., with 5 species from South America.

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Types of Weed 107

 Melissia Hook. f., monotypic genus from Santa Elena with the common
name St. Elena boxwood.

 Physalisastrum Makino, with 9 Asiatic species.

 Tubocapsicum (Wettst.) Makino, with just one species endemic to China.

 Withania Pauq., with 10 species native to the Canary Islands, Africa and Nepal.

 Cuatresia Hunz., with 11 neotropical species. Molecular studies indicate


that this genus, along with Deprea and Larnax has an uncertain taxonomic
position.

 Deprea Raf., with 6 neotropical species.

 Larnax Miers, many taxonomists consider it to be a synonym for Deprea,


contains 22 species native to the Andes.

• Tribe Solaneae. The genera Cyphomandra Sendtn., Discopodium Hochst. , Normania


Lowe, Triguera Cav. and Lycopersicum Mill have been transferred to Solanum. The sub-
tribe is therefore composed of two genera:

 Jaltomata Schltdl., which contains 50 neotropical species.

 Solanum L., the largest genus in the family and one of the broadest of the
angiosperms, with 1,328 species distributed across the whole world.

• Genera with doubtful taxonomic positions (Incertae sedis)

The following genera have still not been placed in any of the recognized subfamilies within the
solanaceas.

• Duckeodendron Kuhlmannb, monotypic genus from the Amazon rainforest.

• Parabouchetia Baillon, poorly-known,monotypic genus from Brazil.

• Pauia Deb. & Dutta, monotypic genus from Assam and Arunachal Pradesh in N.E.India

Genera and Distribution of Species

Flowers and foliage of Cestrum parqui.

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108 Fundamentals of Weed Science

The Solanaceae contain 98 genera and some 2,700 species. Despite this immense richness of spe-
cies, they are not uniformly distributed between the genera. The eight most important genera con-
tain more than 60% of the species, as shown in the table below. Solanum – the genus that typifies
the family - includes nearly 50% of the total species of the solanaceas.

Genera Approximate number of species


Solanum 1,330
Lycianthes 200
Cestrum 150
Nolana 89
Physalis 85
Lycium 85
Nicotiana 76
Brunfelsia 45
Estimated number of species in the family 2,700

Economic Importance
The solanaceas include such important food species as the potato (Solanum tuberosum), the to-
mato (Solanum lycopersicum), the pepper (Capsicum annuum) and the aubergine or egg plant
(Solanum melongena). Nicotiana tabacum, originally from South America, is now cultivated
throughout the world to produce tobacco. Many solanaceas are important weeds in various parts
of the world. Their importance lies in the fact that they can host pathogens or diseases of the cul-
tivated plants, therefore their presence increases the loss of yield or the quality of the harvested
product. An example of this can be seen with Acnistus arborescens and Browalia americana that
host thrips, which cause damage to associated cultivated plants, and certain species of Datura that
play host to various types of virus that are later transmitted to cultivated solanaceas. Some species
of weeds such as, for example Solanum mauritianum in South Africa represent such serious eco-
logical and economic problems that studies are being carried out with the objective of developing
a biological control through the use of insects.

Pink flower of the Brugmansia

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Types of Weed 109

Petunia hybrida, a herbaceous annual that is commonly used in gardens

Various solanaceas species are grown as ornamental trees or shrubs. Examples include Brugman-
sia x candida (“Angel’s Trumpet”) grown for its large pendulous trumpet-shaped flowers, or Brun-
felsia latifolia, whose flowers are very fragrant and change colour from violet to white over a period
of 3 days. Other shrub species that are grown for their attractive flowers are Lycianthes ranton-
netii (Blue Potato Bush or Paraguay Nightshade) with violet-blue flowers and Nicotiana glauca
(“Tree Tobacco”) Other solanacea species and genera that are grown as ornamentals are the petu-
nia (Petunia × hybrida), Lycium, Solanum, Cestrum, Calibrachoa × hybrida and Solandra. There
is even a hybrid between Petunia and Calibrachoa (which constitutes a new nothogenus called ×
Petchoa G. Boker & J. Shaw) that is being sold as an ornamental. Many other species, in particular
those that produce alkaloids, are used in pharmacology and medicine (Nicotiana, Hyoscyamus,
and Datura).

Solanaceas and the Genome


Many of the species belonging to this family, among them tobacco and the tomato, are model
organisms that are used for research into fundamental biological questions. One of the aspects of
the solanaceas’ genomics is an international project that is trying to understand how the same col-
lection of genes and proteins can give rise to a group of organisms that are so morphologically and
ecologically different. The first objective of this project was to sequence the genome of the tomato.
In order to achieve this each of the 12 chromosomes of the tomato’s haploid genome was assigned
to different sequencing centres in different countries. So chromosomes 1 and 10 were sequenced in
the United States, 3 and 11 in China, 2 in Korea, 4 in Britain, 5 in India, 7 in France, 8 in Japan, 9
in Spain and 12 in Italy. The sequencing of the mitochondrial genome was carried out in Argentina
and the chloroplast genome was sequenced in the European Union.

References
• Robert L. Zimdahl (12 March 2004). Weed-crop competition: a review. Wiley-Blackwell. ISBN 978-0-8138-
0279-4. Retrieved 31 July 2010.

• Wagstaff, D. Jesse (2008). International Poisonous Plants Checklist: An Evidence-Based Reference. CRC Press.
p. 1. ISBN 1420062522. Retrieved October 7, 2012.

• Bisby, Frank (1994). Phytochemical Dictionary of the Leguminosae, Volume 1. CRC Press. p. 1. ISBN
0412397706. Retrieved October 7, 2012.

• Knight, Anthony; Walter, Richard (2001). A Guide to Plant Poisoning of Animals in North America. Teton
NewMedia. p. 121. ISBN 1893441113. Retrieved October 7, 2012.

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110 Fundamentals of Weed Science

• Harrison, Lorraine (2012). RHS Latin for gardeners. United Kingdom: Mitchell Beazley. p. 224. ISBN 978-1-
84533-731-5.

• RHS A-Z encyclopedia of garden plants. United Kingdom: Dorling Kindersley. 2008. p. 1136. ISBN 1-4053-
3296-4.

• Gerard, John; Woodward, Marcus (ed.) (1985). Gerard’s Herbal: The History of Plants. New York: Crescent
Books. ISBN 0-517-46470-5

• Howell, Clayson (May 2008). Consolidated list of environmental weeds in New Zealand (PDF). DRDS292.
Wellington: Department of Conservation. ISBN 978-0-478-14413-0. Retrieved 2009-05-06.

• Shurtleff, William; Aoyagi, Akiko (1977). The Book of Kudzu: A Culinary & Healing Guide. Soyinfo Center. p. 9.
ISBN 978-0-394-42068-4. Retrieved 25 October 2013.

• Marchese, C. Marina; Flottum, Kim (2013). The Honey Connoisseur. Black Dog & Leventhal. ISBN
9781603763325.

• William Shurtleff; Akiko Aoyagi (1977). The book of kudzu: a culinary & healing guide. Soyinfo Center. ISBN
9780394420684.

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3
Species of Weed: Invasive and Introduced
Invasive species are species such as plants, fungi or even animals that are not native to a particular
location. Chrysanthemoides monilifera, Heracleum mantegazzianum, Fallopia japonica, Pilosella
aurantiaca and Asparagus asparagoides are some of the examples given in this chapter. The chap-
ter strategically encompasses and incorporates the main species of weed, providing a complete
understanding.

Invasive Species

Beavers from North America constitute an invasive species in Tierra del Fuego, where they have a substantial impact
on landscape and local ecology through their dams.

An invasive species is a plant, fungus, or animal species that is not native to a specific location (an
introduced species), and which has a tendency to spread to a degree believed to cause damage to
the environment, human economy or human health.

One study pointed out widely divergent perceptions of the criteria for invasive species among re-
searchers (p. 135) and concerns with the subjectivity of the term “invasive” (p. 136). Some of the
alternate usages of the term are below:

• The term as most often used applies to introduced species (also called “non-indigenous” or
“non-native”) that adversely affect the habitats and bioregions they invade economically,
environmentally, or ecologically. Such invasive species may be either plants or animals and
may disrupt by dominating a region, wilderness areas, particular habitats, or wildland–
urban interface land from loss of natural controls (such as predators or herbivores). This
includes non-native invasive plant species labeled as exotic pest plants and invasive exotics
growing in native plant communities. It has been used in this sense by government orga-
nizations as well as conservation groups such as the International Union for Conservation
of Nature (IUCN) and the California Native Plant Society. The European Union defines

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112 Fundamentals of Weed Science

“Invasive Alien Species” as those that are, firstly, outside their natural distribution area,
and secondly, threaten biological diversity. It is also used by land managers, botanists, re-
searchers, horticulturalists, conservationists, and the public for noxious weeds. The kudzu
vine (Pueraria lobata), Andean Pampas grass (Cortaderia jubata), and yellow starthistle
(Centaurea solstitialis) are examples.

• An alternate usage broadens the term to include indigenous or “native” species along with
non-native species, that have colonized natural areas (p. 136). Deer are an example, con-
sidered to be overpopulating their native zones and adjacent suburban gardens, by some in
the Northeastern and Pacific Coast regions of the United States.

• Sometimes the term is used to describe a non-native or introduced species that has become
widespread (p. 136). However, not every introduced species has adverse effects on the en-
vironment. A nonadverse example is the common goldfish (Carassius auratus), which is
found throughout the United States, but rarely achieves high densities (p. 136).

Kudzu, a Japanese vine species invasive in the southeast United States, growing in Atlanta, Georgia

Causes
Scientists include species- and ecosystem factors among the mechanisms that when combined,
establish invasiveness in a newly introduced species.

Species-based Mechanisms
While all species compete to survive, invasive species appear to have specific traits or specific combi-
nations of traits that allow them to outcompete native species. In some cases, the competition is about
rates of growth and reproduction. In other cases, species interact with each other more directly.

Researchers disagree about the usefulness of traits as invasiveness markers. One study found that
of a list of invasive and noninvasive species, 86% of the invasive species could be identified from
the traits alone. Another study found invasive species tended to have only a small subset of the
presumed traits and that many similar traits were found in noninvasive species, requiring other
explanations. Common invasive species traits include the following:

• Fast growth

• Rapid reproduction

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Species of Weed: Invasive and Introduced 113

• High dispersal ability

• Phenotypic plasticity (the ability to alter growth form to suit current conditions)

• Tolerance of a wide range of environmental conditions (Ecological competence)

• Ability to live off of a wide range of food types (generalist)

• Association with humans

• Prior successful invasions

Typically, an introduced species must survive at low population densities before it becomes inva-
sive in a new location. At low population densities, it can be difficult for the introduced species to
reproduce and maintain itself in a new location, so a species might reach a location multiple times
before it becomes established. Repeated patterns of human movement, such as ships sailing to and
from ports or cars driving up and down highways offer repeated opportunities for establishment
(also known as a high propagule pressure).

An introduced species might become invasive if it can outcompete native species for resources such
as nutrients, light, physical space, water, or food. If these species evolved under great competition
or predation, then the new environment may host fewer able competitors, allowing the invader to
proliferate quickly. Ecosystems in which are being used to their fullest capacity by native species
can be modeled as zero-sum systems in which any gain for the invader is a loss for the native.
However, such unilateral competitive superiority (and extinction of native species with increased
populations of the invader) is not the rule. Invasive species often coexist with native species for
an extended time, and gradually, the superior competitive ability of an invasive species becomes
apparent as its population grows larger and denser and it adapts to its new location.

Lantana growing in abandoned citrus plantation; Moshav Sdei Hemed, Israel

An invasive species might be able to use resources that were previously unavailable to native spe-
cies, such as deep water sources accessed by a long taproot, or an ability to live on previously un-
inhabited soil types. For example, barbed goatgrass (Aegilops triuncialis) was introduced to Cali-
fornia on serpentine soils, which have low water-retention, low nutrient levels, a high magnesium/
calcium ratio, and possible heavy metal toxicity. Plant populations on these soils tend to show low
density, but goatgrass can form dense stands on these soils and crowd out native species that have
adapted poorly to serpentine soils.

Invasive species might alters its environment by releasing chemical compounds, modifying abiotic
factors, or affecting the behaviour of herbivores, creating a positive or negative impact on oth-
er species. Some species, like Kalanchoe daigremontana, produce allelopathic compounds, that

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114 Fundamentals of Weed Science

might have an inhibitory effect on competing species. Other species like Stapelia gigantea facili-
tates the recruitment of seedlings of other species in arid environments by providing appropriate
microclimatic conditions and preventing herbivory in early stages of development.

Another examples are Centaurea solstitialis (yellow starthistle) and Centaurea diffusa (diffuse
knapweed). These Eastern European noxious weeds have spread through the western and West
Coast states. Experiments show that 8-hydroxyquinoline, a chemical produced at the root of C.
diffusa, has a negative effect only on plants that have not co-evolved with it. Such co-evolved na-
tive plants have also evolved defenses. C. diffusa and C. solstitialis do not appear in their native
habitats to be overwhelmingly successful competitors. Success or lack of success in one habitat
does not necessarily imply success in others. Conversely, examining habitats in which a species is
less successful can reveal novel weapons to defeat invasiveness.

Changes in fire regimens are another form of facilitation. Bromus tectorum, originally from Eur-
asia, is highly fire-adapted. It not only spreads rapidly after burning but also increases the frequen-
cy and intensity (heat) of fires by providing large amounts of dry detritus during the fire season
in western North America. In areas where it is widespread, it has altered the local fire regimen so
much that native plants cannot survive the frequent fires, allowing B. tectorum to further extend
and maintain dominance in its introduced range.

Facilitation also occurs where one species physically modifies a habitat in ways that are advan-
tageous to other species. For example, zebra mussels increase habitat complexity on lake floors,
providing crevices in which invertebrates live. This increase in complexity, together with the nu-
trition provided by the waste products of mussel filter-feeding, increases the density and diversity
of benthic invertebrate communities.

Ecosystem-based Mechanisms
In ecosystems, the amount of available resources and the extent to which those resources are used
by organisms determines the effects of additional species on the ecosystem. In stable ecosystems,
equilibrium exists in the use of available resources. These mechanisms describe a situation in which
the ecosystem has suffered a disturbance, which changes the fundamental nature of the ecosystem.

When changes such as a forest fire occur, normal succession favors native grasses and forbs. An
introduced species that can spread faster than natives can use resources that would have been
available to native species, squeezing them out. Nitrogen and phosphorus are often the limiting
factors in these situations.

Every species occupies a niche in its native ecosystem; some species fill large and varied roles,
while others are highly specialized. Some invading species fill niches that are not used by native
species, and they also can create new niches.An example of this type can be found within the Lam-
propholis delicata species of skink.

Ecosystem changes can alter species’ distributions. For example, edge effects describe what hap-
pens when part of an ecosystem is disturbed as when land is cleared for agriculture. The boundary
between remaining undisturbed habitat and the newly cleared land itself forms a distinct habitat,
creating new winners and losers and possibly hosting species that would not thrive outside the
boundary habitat.

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Species of Weed: Invasive and Introduced 115

One interesting finding in studies of invasive species has shown that introduced populations
have great potential for rapid adaptation and this is used to explain how so many introduced
species are able to establish and become invasive in new environments. When bottlenecks and
founder effects cause a great decrease in the population size and may constrict genetic vari-
ation, the individuals begin to show additive variance as opposed to epistatic variance. This
conversion can actually lead to increased variance in the founding populations which then
allows for rapid adaptive evolution. Following invasion events, selection may initially act on
the capacity to disperse as well as physiological tolerance to the new stressors in the environ-
ment. Adaptation then proceeds to respond to the selective pressures of the new environment.
These responses would most likely be due to temperature and climate change, or the presence
of native species whether it be predator or prey. Adaptations include changes in morphology,
physiology, phenology, and plasticity.

Rapid adaptive evolution in these species leads to offspring that have higher fitness and are better
suited for their environment. Intraspecific phenotypic plasticity, pre-adaptation and post-intro-
duction evolution are all major factors in adaptive evolution. Plasticity in populations allows room
for changes to better suit the individual in its environment. This is key in adaptive evolution be-
cause the main goal is how to best be suited to the ecosystem that the species has been introduced.
The ability to accomplish this as quickly as possible will lead to a population with a very high
fitness. Pre-adaptations and evolution after the initial introduction also play a role in the success
of the introduced species. If the species has adapted to a similar ecosystem or contains traits that
happen to be well suited to the area that it is introduced, it is more likely to fare better in the new
environment. This, in addition to evolution that takes place after introduction, all determine if the
species will be able to become established in the new ecosystem and if it will reproduce and thrive.

Ecology
Traits of Invaded Ecosystems
In 1958, Charles S. Elton claimed that ecosystems with higher species diversity were less subject to
invasive species because of fewer available niches. Other ecologists later pointed to highly diverse,
but heavily invaded ecosystems and argued that ecosystems with high species diversity were more
susceptible to invasion.

This debate hinged on the spatial scale at which invasion studies were performed, and the issue
of how diversity affects susceptibility remained unresolved as of 2011. Small-scale studies tended
to show a negative relationship between diversity and invasion, while large-scale studies tended
to show the reverse. The latter result may be a side-effect of invasives’ ability to capitalize on in-
creased resource availability and weaker species interactions that are more common when larger
samples are considered.

Invasion was more likely in ecosystems that were similar to the one in which the potential invader
evolved. Island ecosystems may be more prone to invasion because their species faced few strong
competitors and predators, or because their distance from colonizing species populations makes them
more likely to have “open” niches. An example of this phenomenon was the decimation of native bird
populations on Guam by the invasive brown tree snake.Conversely, invaded ecosystems may lack the
natural competitors and predators that check invasives’ growth in their native ecosystems.

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116 Fundamentals of Weed Science

The brown tree snake (Boiga irregularis)

Invaded ecosystems may have experienced disturbance, typically human-induced. Such a distur-
bance may give invasive species a chance to establish themselves with less competition from na-
tives less able to adapt to a disturbed ecosystem.

Vectors
Non-native species have many vectors, including biogenic vectors, but most invasions are associ-
ated with human activity. Natural range extensions are common in many species, but the rate and
magnitude of human-mediated extensions in these species tend to be much larger than natural
extensions, and humans typically carry specimens greater distances than natural forces.

An early human vector occurred when prehistoric humans introduced the Pacific rat (Rattus exu-
lans) to Polynesia.

Chinese mitten crab (Eriocheir sinensis)

Vectors include plants or seeds imported for horticulture. The pet trade moves animals across
borders, where they can escape and become invasive. Organisms stow away on transport vehicles.
Ballast water taken up at sea and released in port by transoceanic vessels is the largest vector for
non-native aquatic species invasions. Around the world on the average day, more than 3,000 dif-
ferent species of aquatic life may be transported on these vessels. For example, freshwater zebra
mussels, native to the Black, Caspian and Azov seas, probably reached the Great Lakes via ballast
water from a transoceanic vessel. Although the zebra mussel invasion was first noted in 1988, and
a mitigation plan was successfully implemented shortly thereafter, the plan had (and continued to

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Species of Weed: Invasive and Introduced 117

have as of 2005) a serious flaw or loophole, whereby ships that are loaded with cargo when they
reach the Seaway need not be tested, but all the same they transfer ballast ‘puddles’ between Sea-
way ports.

The arrival of invasive propagules to a new site is a function of the site’s invasibility.

Species have also been introduced intentionally. For example, to feel more “at home,” American
colonists formed “Acclimation Societies” that repeatedly imported birds that were native to Europe
to North America and other distant lands. In 2008, U.S. postal workers in Pennsylvania noticed
noises coming from inside a box from Taiwan; the box contained more than two dozen live beetles.
Agricultural Research Service entomologists identified them as rhinoceros beetle, hercules beetle,
and king stag beetle. Because these species were not native to the U.S., they could have threatened
native ecosystems. To prevent exotic species from becoming a problem in the U.S., special han-
dling and permits are required when living materials are shipped from foreign countries. USDA
programs such as Smuggling Interdiction and Trade Compliance (SITC) attempt to prevent exotic
species outbreaks in America.

Economics plays a major role in exotic species introduction. High demand for the valuable Chinese
mitten crab is one explanation for the possible intentional release of the species in foreign waters.

Impacts of Wildfire
Invasive species often exploit disturbances to an ecosystem (wildfires, roads, foot trails) to colo-
nize an area. Large wildfires can sterilize soils, while adding a variety of nutrients. In the resulting
free-for-all, formerly entrenched species lose their advantage, leaving more room for invasives. In
such circumstances plants that can regenerate from their roots have an advantage. Non-natives
with this ability can benefit from a low intensity fire burns that removes surface vegetation, leav-
ing natives that rely on seeds for propagation to find their niches occupied when their seeds finally
sprout.

Impact of Wildfire Suppression on Spreading


Wildfires often occur in remote areas, needing fire suppression crews to travel through pristine
forest to reach the site. The crews can bring invasive seeds with them. If any of these stowaway
seeds become established, a thriving colony of invasives can erupt in as few as six weeks, after
which controlling the outbreak can need years of continued attention to prevent further spread.
Also, disturbing the soil surface, such as cutting firebreaks, destroys native cover, exposes soil, and
can accelerate invasions. In suburban and wildland-urban interface areas, the vegetation clear-
ance and brush removal ordinances of municipalities for defensible space can result in excessive
removal of native shrubs and perennials that exposes the soil to more light and less competition
for invasive plant species.

Fire suppression vehicles are often major culprits in such outbreaks, as the vehicles are often driv-
en on back roads often overgrown with invasive plant species. The undercarriage of the vehicle
becomes a prime vessel of transport. In response, on large fires, washing stations “decontaminate”
vehicles before engaging in suppression activities.Large wildfires attract firefighters from remote
places, further increasing the potential for seed transport.

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118 Fundamentals of Weed Science

Effects

An American alligator attacking a Burmese python in Florida; the Burmese python is an invasive species which is
posing a threat to many indigenous species, including the alligator

Ecological
Land clearing and human habitation put significant pressure on local species. Disturbed habitats
are prone to invasions that can have adverse effects on local ecosystems, changing ecosystem func-
tions. A species of wetland plant known as ʻaeʻae in Hawaii (the indigenous Bacopa monnieri) is
regarded as a pest species in artificially manipulated water bird refuges because it quickly covers
shallow mudflats established for endangered Hawaiian stilt (Himantopus mexicanus knudseni),
making these undesirable feeding areas for the birds.

Multiple successive introductions of different non-native species can have interactive effects; the
introduction of a second non-native species can enable the first invasive species to flourish. Exam-
ples of this are the introductions of the amethyst gem clam (Gemma gemma) and the European
green crab (Carcinus maenas). The gem clam was introduced into California’s Bodega Harbor
from the East Coast of the United States a century ago. It had been found in small quantities in
the harbor but had never displaced the native clam species (Nutricola spp.). In the mid-1990s, the
introduction of the European green crab, found to prey preferentially on the native clams, resulted
in a decline of the native clams and an increase of the introduced clam populations.

In the Waterberg region of South Africa, cattle grazing over the past six centuries has allowed invasive
scrub and small trees to displace much of the original grassland, resulting in a massive reduction in
forage for native bovids and other grazers. Since the 1970s, large scale efforts have been underway to
reduce invasive species; partial success has led to re-establishment of many species that had dwindled
or left the region. Examples of these species are giraffe, blue wildebeest, impala, kudu and white rhino.

Invasive species can change the functions of ecosystems. For example, invasive plants can alter the
fire regimen (cheatgrass, Bromus tectorum), nutrient cycling (smooth cordgrass Spartina alterni-
flora), and hydrology (Tamarix) in native ecosystems. Invasive species that are closely related to
rare native species have the potential to hybridize with the native species. Harmful effects of hy-
bridization have led to a decline and even extinction of native species. For example, hybridization
with introduced cordgrass, Spartina alterniflora, threatens the existence of California cordgrass
(Spartina foliosa) in San Francisco Bay. Invasive species cause competition for native species and
because of this 400 of the 958 endangered species under the Endangered Species Act are at risk

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Species of Weed: Invasive and Introduced 119

Geomorphological
Primary geomorphological effects of invasive plants are bioconstruction and bioprotection.
For example, Kudzu Pueraria montana, a vine native to Asia was widely introduced in the
southeastern USA in the early 20th century to control soil erosion. While primary effects of
invasive animals are bioturbation, bioerosion, and bioconstruction. For example, invasion of
Chinese mitten crab Eriocheir sinensis have resulted in higher bioturbation and bioerosion
rates.

Economic
Economic Opportunities
Some invasions offer potential commercial benefits. For instance, silver carp and common carp
can be harvested for human food and exported to markets already familiar with the product, or
processed into pet foods, or mink feed. Water hyacinth can be turned into fuel by methane digest-
ers, and other invasive plants can also be harvested and utilized as a source of bioenergy.

Benefits
Non-native species can have benefits. Asian oysters, for example, filter water pollutants better
than native oysters. They also grow faster and withstand disease better than natives. Biologists are
currently considering releasing this mollusk in the Chesapeake Bay to help restore oyster stocks
and remove pollution. A recent study by the Johns Hopkins School of Public Health found the
Asian oyster could significantly benefit the bay’s deteriorating water quality. Additionally, some
species have invaded an area so long ago that they have found their own beneficial niche in the
environment. For example, L. leucozonium, shown by population genetic analysis to be an invasive
species in North America, has become an important pollinator of caneberry as well as cucurbit,
apple trees, and blueberry bushes.

Invasivorism
Invasive species are flora and fauna whose introduction into a habitat disrupts the native eco-sys-
tem. In response, Invasivorism is a movement that explores the idea of eating invasive species in
order to control, reduce, or eliminate their populations. Chefs from around the world have begun
seeking out and using invasive species as alternative ingredients. Miya’s of New Haven, Connecti-
cut created the first invasive species menu in the world. Skeptics point out that once a foreign
species has entrenched itself in a new place—such as the Indo-Pacific lionfish that has now virtu-
ally taken over the waters of the Western Atlantic, Caribbean and Gulf of Mexico—eradication is
almost impossible. Critics argue that encouraging consumption might have the unintended effect
of spreading harmful species even more widely.

Proponents of invasivorism argue that humans have the ability to eat away any species that it has
an appetite for, pointing to the many animals which humans have been able to hunt to extinction
- such as the Dodo bird, the Caribbean monk seal, and the Passenger pigeon. Proponents of inva-
sivorism also point to the success that Jamaica has had in significantly decreasing the population
of lionfish by encouraging the consumption of the fish.

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120 Fundamentals of Weed Science

A dish that features whole fried invasive lionfish at Fish Fish of Miami, Florida

Costs
Economic costs from invasive species can be separated into direct costs through production loss
in agriculture and forestry, and management costs. Estimated damage and control cost of invasive
species in the U.S. alone amount to more than $138 billion annually. Economic losses can also
occur through loss of recreational and tourism revenues. When economic costs of invasions are
calculated as production loss and management costs, they are low because they do not consider
environmental damage; if monetary values were assigned to the extinction of species, loss in bio-
diversity, and loss of ecosystem services, costs from impacts of invasive species would drastically
increase. The following examples from different sectors of the economy demonstrate the impact of
biological invasions.

Plant Industry
Weeds reduce yield in agriculture, though they may provide essential nutrients. Some deep-rooted
weeds can “mine” nutrients from the subsoil and deposit them on the topsoil, while others provide
habitat for beneficial insects or provide foods for pest species. Many weed species are accidental
introductions that accompany seeds and imported plant material. Many introduced weeds in pas-
tures compete with native forage plants, threaten young cattle (e.g., leafy spurge, Euphorbia esula)
or are unpalatable because of thorns and spines (e.g., yellow starthistle). Forage loss from invasive
weeds on pastures amounts to nearly US$1 billion in the U.S. alone. A decline in pollinator services
and loss of fruit production has been caused by honey bees infected by the invasive varroa mite.
Introduced rats (Rattus rattus and R. norvegicus) have become serious pests on farms, destroying
stored grains.

Invasive plant pathogens and insect vectors for plant diseases can also suppress agricultural yields
and nursery stock. Citrus greening is a bacterial disease vectored by the invasive Asian citrus psyl-
lid (ACP). Because of the impacts of this disease on citrus crops, citrus is under quarantine and
highly regulated in areas where ACP has been found.

Aquaculture
Aquaculture is a very common vector of species introductions – mainly of species with economic
potential (e.g., Oreochromis niloticus)

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Species of Weed: Invasive and Introduced 121

Forestry

Poster asking campers to not move firewood around, avoiding the spread of invasive species.

The unintentional introduction of forest pest species and plant pathogens can change forest ecol-
ogy and damage the timber industry. Overall, forest ecosystems in the U.S. are widely invaded by
exotic pests, plants, and pathogens.

The Asian long-horned beetle (Anoplophora glabripennis) was first introduced into the U.S. in
1996, and was expected to infect and damage millions of acres of hardwood trees. As of 2005
thirty million dollars had been spent in attempts to eradicate this pest and protect millions of
trees in the affected regions. The woolly adelgid has inflicted damage on old-growth spruce, fir
and hemlock forests and damages the Christmas tree industry. And the chestnut blight fun-
gus (Cryphonectria parasitica) and Dutch elm disease (Ophiostoma novo-ulmi) are two plant
pathogens with serious impacts on these two species, and forest health. Garlic mustard, Alliaria
petiolata, is one of the most problematic invasive plant species in eastern North American for-
ests. The characteristics of garlic mustard are slightly different from those of the surrounding
native plants, which results in a highly successful species that is altering the composition and
function of the native communities it invades. When garlic mustard invades the understory of
a forest, it affects the growth rate of tree seedlings, which is likely to alter forest regeneration of
impact forest composition in the future.

Tourism and Recreation


Invasive species can impact outdoor recreation, such as fishing, hunting, hiking, wildlife viewing,
and water-based activities. They can damage a wide array of environmental services that are im-
portant to recreation, including, but not limited to, water quality and quantity, plant and animal
diversity, and species abundance. Eiswerth states, “very little research has been performed to esti-
mate the corresponding economic losses at spatial scales such as regions, states, and watersheds.”
Eurasian watermilfoil (Myriophyllum spicatum) in parts of the US, fill lakes with plants com-
plicating fishing and boating. The very loud call of the introduced common coqui depresses real
estate values in affected neighborhoods of Hawaii.

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122 Fundamentals of Weed Science

Health
Encroachment of humans into previously remote ecosystems has exposed exotic diseases such as
HIV to the wider population. Introduced birds (e.g. pigeons), rodents and insects (e.g. mosquito,
flea, louse and tsetse fly pests) can serve as vectors and reservoirs of human afflictions. The intro-
duced Chinese mitten crabs are carriers of Asian lung fluke. Throughout recorded history, epidem-
ics of human diseases, such as malaria, yellow fever, typhus, and bubonic plague, spread via these
vectors. A recent example of an introduced disease is the spread of the West Nile virus, which killed
humans, birds, mammals, and reptiles. Waterborne disease agents, such as cholera bacteria (Vibrio
cholerae), and causative agents of harmful algal blooms are often transported via ballast water. In-
vasive species and accompanying control efforts can have long term public health implications. For
instance, pesticides applied to treat a particular pest species could pollute soil and surface water.

Biodiversity
Biotic invasion is considered one of the five top drivers for global biodiversity loss and is increasing
because of tourism and globalization.This may be particularly true in inadequately regulated fresh
water systems, though quarantines and ballast water rules have improved the situation.

Invasive species may drive local native species to extinction via competitive exclusion, niche dis-
placement, or hybridisation with related native species. Therefore, besides their economic ramifi-
cations, alien invasions may result in extensive changes in the structure, composition and global
distribution of the biota of sites of introduction, leading ultimately to the homogenisation of the
world’s fauna and flora and the loss of biodiversity. Nevertheless, it is difficult to unequivocally
attribute extinctions to a species invasion, and the few scientific studies that have done so have
been with animal taxa. Concern over the impacts of invasive species on biodiversity must therefore
consider the actual evidence (either ecological or economic), in relation to the potential risk.

Alien invasive species Parthenium hysterophorus smothering native flora in Achanakmar Tiger
Reserve, Bilaspur, Chhattisgarh, India.

Genetic Pollution
Native species can be threatened with extinction through the process of genetic pollution. Ge-
netic pollution is unintentional hybridization and introgression, which leads to homogenization
or replacement of local genotypes as a result of either a numerical or fitness advantage of the

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Species of Weed: Invasive and Introduced 123

introduced species. Genetic pollution can operate either through introduction or through habitat
modification, bringing previously isolated species into contact. Hybrids resulting from rare species
that interbreed with abundant species can swamp the rarer species’ gene pool. This is not always
apparent from morphological observations alone. Some degree of gene flow is normal, and pre-
serves constellations of genes and genotypes. An example of this is the interbreeding of migrating
coyotes with the red wolf, in areas of eastern North Carolina where the red wolf was reintroduced.

Study

Stage Characteristic

0 Propagules residing in a donor region

I Traveling

II Introduced

III Localized and numerically rare

IVa Widespread but rare

IVb Localized but dominant

V Widespread and dominant

While the study of invasive species can be done within many subfields of biology, the majority of
research on invasive organisms has been within the field of ecology and geography where the issue
of biological invasions is especially important. Much of the study of invasive species has been in-
fluenced by Charles Elton’s 1958 book The Ecology of Invasion by Animals and Plants which drew
upon the limited amount of research done within disparate fields to create a generalized picture of
biological invasions. Studies on invasive species remained sparse until the 1990s when research in
the field experienced a large amount of growth which continues to this day. This research, which
has largely consisted of field observational studies, has disproportionately been concerned with
terrestrial plants. The rapid growth of the field has driven a need to standardize the language used
to describe invasive species and events. Despite this, little standard terminology exists within the
study of invasive species which itself lacks any official designation but is commonly referred to as
“Invasion ecology” or more generally “Invasion biology”. This lack of standard terminology is a
significant problem, and has largely arisen due to the interdisciplinary nature of the field which
borrows terms from numerous disciplines such as agriculture, zoology, and pathology, as well as
due to studies on invasive species being commonly performed in isolation of one another.

In an attempt to avoid the ambiguous, subjective, and pejorative vocabulary that so often accom-
panies discussion of invasive species even in scientific papers, Colautti and MacIsaac proposed a
new nomenclature system based on biogeography rather than on taxa.

By discarding taxonomy, human health, and economic factors, this model focused only on ecolog-
ical factors. The model evaluated individual populations rather than entire species. It classified
each population based on its success in that environment. This model applied equally to indige-
nous and to introduced species, and did not automatically categorize successful introductions as
harmful.

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124 Fundamentals of Weed Science

Introduced Species
An introduced, alien, exotic, non-indigenous, or non-native species, or simply an introduction, is
a species living outside its native distributional range, which has arrived there by human activity,
either deliberate or accidental. Non-native species can have various effects on the local ecosystem.
Introduced species that become established and spread beyond the place of introduction are called
invasive species. Some have a negative effect on a local ecosystem. Some introduced species may
have no negative effect or only minor impact. Some species have been introduced intentionally to
combat pests. They are called biocontrols and may be regarded as beneficial as an alternative to
pesticides in agriculture for example. In some instances the potential for being beneficial or detri-
mental in the long run remains unknown.

Sweet clover (Melilotus sp.), introduced and naturalized to the U.S. from Eurasia as a forage and cover crop.

The effects of introduced species on natural environments have gained much scrutiny from scien-
tists, governments, farmers and others.

Terminology
The terminology associated with introduced species is now in flux for various reasons. Other terms
with somewhat similar meanings) with introduced are acclimatized, adventive, naturalized, and
immigrant but those terms refer to a subset of introduced species: those that have become estab-
lished and can reproduce without human assistance. The term invasive refers only to those species
that become established and spread beyond the place of introduction. For practical purposes, this
term is applied only to invasive species that cause damage.

In the broadest and most widely used sense, an introduced species is synonymous with non-native
and therefore applies as well to most garden and farm organisms; these adequately fit the basic
definition given above. However, some sources add to that basic definition “and are now repro-

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Species of Weed: Invasive and Introduced 125

ducing in the wild,” which removes from consideration as introduced all of those species raised or
grown in gardens or farms that do not survive without tending by people. With respect to plants,
these latter are in this case defined as either ornamental or cultivated plants.

The following definition from the United States Environmental Protection Agency (EPA), although
perhaps lacking ecological sophistication, is more typical: introduced species “A species that has
been intentionally or inadvertently brought into a region or area. Also called an exotic or non-na-
tive species.” Introduction of a species outside its native range is all that is required to be qualified
as an “introduced species” such that one can distinguish between introduced species that may not
occur except in cultivation, under domestication or captivity whereas others become established
outside their native range and reproduce without human assistance. Such species might be termed
“naturalized”, “established”, “wild non-native species”. If they further spread beyond the place of
introduction they are called “invasive”. The transition from introduction, to establishment and to
invasion has been described in the context of plants. Introduced species are essentially “non-na-
tive” species. Invasive species are those introduced species that spreadwidely or quickly and cause
harm, be that to the environment, human health, other valued resources or the economy. There
have been calls from scientists to consider a species “invasive” only in terms of their spread and
reproduction rather than the harm they may cause.

According to a practical definition, an invasive species is one that has been introduced and become
a pest in its new location, spreading (invading) by natural means. The term is used to imply both a
sense of urgency and actual or potential harm. For example, U.S. Executive Order 13112 (1999) de-
fines “invasive species” as “an alien species whose introduction does or is likely to cause economic
or environmental harm or harm to human health”. The biological definition of invasive species, on
the other hand, makes no reference to the harm they may cause, only to the fact that they spread
beyond the area of original introduction.

Although some argue that “invasive” is a loaded word and harm is difficult to define, the fact of the
matter is that organisms have and continue to be introduced to areas in which they are not native,
sometimes with but usually without much regard to the harm that could result.

From a regulatory perspective, it is neither desirable nor practical to list as undesirable or outright
ban all non-native species (although the State of Hawaii has adopted an approach that comes
close to this). Regulations require a definitional distinction between non-natives that are deemed
especially onerous and all others. Introduced pest species that are officially listed as invasive, best
fit the definition of an invasive species. Early detection and rapid response is the most effective
strategy for regulating a pest species and reducing economic and environmental impacts of an
introduction

In Great Britain, the Wildlife and Countryside Act 1981 prevents the introduction of any animal
not naturally occurring in the wild or any of a list of both animals or plants introduced previously
and proved to be invasive.

Nature of Introductions
By definition, a species is considered “introduced” when its transport into an area outside of its
native range is human mediated. Introductions by humans can be described as either intentional

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126 Fundamentals of Weed Science

or accidental. Intentional introductions have been motivated by individuals or groups who either
(1) believe that the newly introduced species will be in some way beneficial to humans in its new
location or, (2) as is the case with pythons in the Everglades, species are introduced intentionally
but with no regard to the potential impact. Unintentional or accidental introductions are most of-
ten a byproduct of human movements, and are thus unbound to human motivations. Subsequent
range expansion of introduced species may or may not involve human activity.

Intentional Introductions
Species that humans intentionally transport to new regions can subsequently become successful-
ly established in two ways. In the first case, organisms are purposely released for establishment
in the wild. It is sometimes difficult to predict whether a species will become established upon
release, and if not initially successful, humans have made repeated introductions to improve the
probability that the species will survive and eventually reproduce in the wild. In these cases it is
clear that the introduction is directly facilitated by human desires.

Male Lophura nycthemera (silver pheasant), a native of East Asia that has been introduced into parts of Europe for
ornamental reasons.

In the second case, species intentionally transported into a new region may escape from captive
or cultivated populations and subsequently establish independent breeding populations. Escaped
organisms are included in this category because their initial transport to a new region is human
motivated.

Perhaps the most common motivation for introducing a species into a new place is that of econom-
ic gain. Examples of species introduced for the purposes of benefiting agriculture, aquaculture or
other economic activities are widespread. Eurasian carp was first introduced to the United States
as a potential food source. The apple snail was released in Southeast Asia with the intent that it be
used as a protein source, and subsequently to places like Hawaii to establish a food industry. In
Alaska, foxes were introduced to many islands to create new populations for the fur trade. About
twenty species of African and European dung beetles have established themselves in Australia
after deliberate introduction by the Australian Dung Beetle Project in an effort to reduce the im-
pact of livestock manure. The timber industry promoted the introduction of Monterey pine (Pinus
radiata) from California to Australia and New Zealand as a commercial timber crop. These exam-
ples represent only a small subsample of species that have been moved by humans for economic
interests.

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Species of Weed: Invasive and Introduced 127

Introductions have also been important in supporting recreation activities or otherwise increasing
human enjoyment. Numerous fish and game animals have been introduced for the purposes of
sport fishing and hunting (Earthworms as invasive species). The introduced amphibian (Ambysto-
ma tigrinum) that threatens the endemic California salamander (Ambystoma californiense) was
introduced to California as a source of bait for fishermen. Pet animals have also been frequently
transported into new areas by humans, and their escapes have resulted in several successful intro-
ductions, such as those of feral cats and parrots.

Many plants have been introduced with the intent of aesthetically improving public recreation ar-
eas or private properties. The introduced Norway maple for example occupies a prominent status
in many of Canada’s parks. The transport of ornamental plants for landscaping use has and contin-
ues to be a source of many introductions. Some of these species have escaped horticultural control
and become invasive. Notable examples include water hyacinth, salt cedar, and purple loosestrife.

In other cases, species have been translocated for reasons of “cultural nostalgia,” which refers to
instances in which humans who have migrated to new regions have intentionally brought with
them familiar organisms. Famous examples include the introduction of starlings to North America
by Englishman Eugene Schieffelin, a lover of the works of Shakespeare and the chairman of the
American Acclimatization Society, who, it is rumoured, wanted to introduce all of the birds men-
tioned in Shakespeare’s plays into the United States. He deliberately released eighty starlings into
Central Park in New York City in 1890, and another forty in 1891.

Yet another prominent example is the introduction of the European rabbit to Australia by one
Thomas Austin, a British landowner who had the rabbits released on his estate in Victoria because
he missed hunting them. A more recent example is the introduction of the common wall lizard to
North America by a Cincinnati boy, George Rau, around 1950 after a family vacation to Italy.

Intentional introductions have also been undertaken with the aim of ameliorating environmental
problems. A number of fast spreading plants such as garlic mustard and kudzu have been intro-
duced as a means of erosion control. Other species have been introduced as biological control
agents to control invasive species and involves the purposeful introduction of a natural enemy of
the target species with the intention of reducing its numbers or controlling its spread.

A special case of introduction is the reintroduction of a species that has become locally endangered
or extinct, done in the interests of conservation. Examples of successful reintroductions include
wolves to Yellowstone National Park in the U.S., and the red kite to parts of England and Scot-
land. Introductions or translocations of species have also been proposed in the interest of genetic
conservation, which advocates the introduction of new individuals into genetically depauperate
populations of endangered or threatened species.

The above examples highlight the intent of humans to introduce species as a means of incurring
some benefit. While these benefits have in some cases been realized, introductions have also resulted
in unforeseen costs, particularly when introduced species take on characteristics of invasive species.

Non-native species can become such a common part of an environment, culture, and even diet
that little thought is given to their geographic origin. For example, soybeans, kiwi fruit, wheat and
all livestock except the American bison and the turkey are non-native species to North America.
Collectively, non-native crops and livestock comprise 98% of US food. These and other benefits

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128 Fundamentals of Weed Science

from non-natives are so vast that, according to the Congressional Research Service, they probably
exceed the costs.

Unintentional Introductions
Unintentional introductions occur when species are transported by human vectors. For example, three
species of rat (the black, Norway and Polynesian) have spread to most of the world as hitchhikers
on ships. There are also numerous examples of marine organisms being transported in ballast water,
one being the zebra mussel. Over 200 species have been introduced to the San Francisco Bay in this
manner making it the most heavily invaded estuary in the world. Increasing rates of human travel are
providing accelerating opportunities for species to be accidentally transported into areas in which they
are not considered native. There is also the accidental release of the Africanized honey bees (AHB),
known colloquially as “killer bees” or Africanized bee to Brazil in 1957 and the Asian carps to the United
States. The insect commonly known as the brown marmorated stink bug (Halyomorpha halys) was
introduced accidentally in Pennsylvania. Another form of unintentional introductions is when an in-
tentionally introduced plant carries a parasite or herbivore with it. Some become invasive, for example
the oleander aphid, accidentally introduced with the ornamental plant, oleander.

Introduced Plants
Many non-native plants have been introduced into new territories, initially as either ornamental
plants or for erosion control, stock feed, or forestry. Whether an exotic will become an invasive
species is seldom understood in the beginning, and many non-native ornamentals languish in the
trade for years before suddenly naturalizing and becoming invasive.

Peaches, for example, originated in China, and have been carried to much of the populated world.
Tomatoes are native to the Andes. Squash (pumpkins), maize (corn), and tobacco are native to
the Americas, but were introduced to the Old World. Many introduced species require continued
human intervention to survive in the new environment. Others may become feral, but do not seri-
ously compete with natives, but simply increase the biodiversity of the area.

Dandelions are also introduced species to North America.

A very troublesome marine species in southern Europe is the seaweed Caulerpa taxifolia. Caul-
erpa was first observed in the Mediterranean Sea in 1984, off the coast of Monaco. By 1997, it had
covered some 50 km². It has a strong potential to overgrow natural biotopes, and represents a
major risk for sublittoral ecosystems. The origin of the alga in the Mediterranean was thought to
be either as a migration through the Suez Canal from the Red Sea, or as an accidental introduction
from an aquarium. Another troublesome plant species is the terrestrial plant Phyla canescens,
which was intentionally introduced into many countries in North America, Europe, and Africa
as an ornamental plant. This species has become invasive in Australia, where it threatens native
rare plants and causes erosion and soil slumping around river banks. It has also become invasive
in France where it has been listed as an invasive plant species of concern in the Mediterranean
region, where it can form monocultures that threaten critical conservation habitats.

Japanese knotweed grows profusely in many nations. Human beings introduced it into many plac-
es in the 19th century. It is a source of resveratrol, a dietary supplement.

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Species of Weed: Invasive and Introduced 129

Introduced Animals
Bear in mind that most introduced species do not become invasive. Examples of introduced ani-
mals that have become invasive include the gypsy moth in eastern North America, the zebra mus-
sel and alewife in the Great Lakes, the Canada goose and gray squirrel in Europe, the muskrat in
Europe and Asia, the cane toad and red fox in Australia, nutria in North America, Eurasia, and
Africa, and the common brushtail possum in New Zealand.

Most Commonly Introduced Species


Some species, such as the brown rat, house sparrow, ring-necked pheasant and European starling,
have been introduced very widely. In addition there are some agricultural and pet species that fre-
quently become feral; these include rabbits, dogs, goats, fish, pigs and cats.

Invasive Exotic Diseases


History is rife with the spread of exotic diseases, such as the introduction of smallpox into the
indigenous peoples of the Americas by the Spanish, where it obliterated entire populations of in-
digenous civilizations before they were ever even seen by Europeans.

Problematic exotic disease introductions in the past century or so include the chestnut blight which
has almost eliminated the American chestnut tree from its forest habitat, and Dutch elm disease,
which has severely reduced the American elm trees in forests and cities.

Diseases may also be vectored by invasive insects such as the Asian citrus psyllid and the bacterial
disease citrus greening.

Introduced Species on Islands


Perhaps the best place to study problems associated with introduced species is on islands. Depend-
ing upon the isolation (how far an island is located from continental biotas), native island biolog-
ical communities may be poorly adapted to the threat posed by exotic introductions. Often this
can mean that no natural predator of an introduced species is present, and the non-native spreads
uncontrollably into open or occupied niche.

An additional problem is that birds native to small islands may have become flightless because of
the absence of predators prior to introductions and cannot readily escape danger. The tendency of
rails in particular to evolve flightless forms on islands has led to the disproportionate number of
extinctions in that family.

The field of island restoration has developed as a field of conservation biology and ecological res-
toration, a great deal of which deals with the eradication of introduced species.

New Zealand
In New Zealand the largest commercial crop is Pinus radiata, the native Californian Monterey
pine tree, which grows as well in New Zealand as in California. However, the pine forests are also
occupied by deer from North America and Europe and by possums from Australia. All are exotic

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130 Fundamentals of Weed Science

species and all have thrived in the New Zealand environment. The pines are seen as beneficial
while the deer and possums are regarded as serious pests.

Common gorse, originally a hedge plant in Britain, was introduced to New Zealand for the same pur-
pose. Like the Monterey pine, it has shown a favour to its new climate. It is, however, regarded as a nox-
ious plant that threatens to obliterate native plants in much of the country and is hence routinely erad-
icated, though it can also provide a nursery environment for native plants to reestablish themselves.

Rabbits, introduced as a food source by sailors in the 1800s, have become a severe nuisance to
farmers, notably in the South Island. The myxomatosis virus was illegally imported and illegally
released, but it had little lasting effect upon the rabbit population other than to make it more re-
sistant to the virus.

Cats, brought by the Māori and later by Europeans, have had a devastating effect upon the native
birdlife, particularly as many New Zealand birds are flightless. Feral cats and dogs which were
originally brought as pets are also known to kill large numbers of birds. A recent (2006) study in
the South Island has shown that even domestic cats with a ready supply of food from their owners
may kill hundreds of birds in a year, including natives.

Sparrows, which were brought to control insects upon the introduced grain crops, have displaced
native birds as have rainbow lorikeets and cockatoos (both from Australia) which fly free around
areas west of Auckland City such as the Waitakere Ranges.

In much of New Zealand, the Australian black swan has effectively eliminated the existence of the
previously introduced mute swan.

Two notable varieties of spiders have also been introduced: the white tail spider and the redback
spider. Both may have arrived inside shipments of fruit. Until then, the only spider (and the only
poisonous animal) dangerous to humans was the native katipo, which is very similar to the red-
back and interbreed with the more aggressive Australian variety.

Introduced Species on a Planetary Body


It has been hypothesized that invasive species of microbial life could contaminate a planetary body
after the former is introduced by a space probe or spacecraft, either deliberately or unintentionally.

Chrysanthemoides Monilifera
Chrysanthemoides monilifera (subspecies of which are known as Boneseed and Bitou Bush in
Australasia, or Bietou, Tick Berry, Bosluisbessie, or Weskusbietou in South Africa) is an evergreen
flowering shrub or small tree, usually with woolly, dull, serrate, oval leaves, but with glossy round
leaves in the subsp. rotundata, belonging to the Asteraceae (daisy) family. It is native to South
Africa, such as the Cape Flats Dune Strandveld habitat.

The plant has become a major environmental weed and invasive species in Australia and New
Zealand.

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Species of Weed: Invasive and Introduced 131

Taxonomy
Chrysanthemoides monilifera is one of only two members of the genus Chrysanthemoides, the
other is Chrysanthemoides incana.

C. monilifera has six recognized subspecies:

• ssp. canescens

• ssp. monilifera

• ssp. pisifera

• ssp. rotundata

• ssp. septentrionalis

• ssp. subcanescens

In Australia, C. m. ssp. monilifera is known by the common name ‘Boneseed’, while C. m. ssp. ro-


tundata is known by the common name ‘Bitou Bush’. In New Zealand subspecies are not distin-
guished and C. monilifera is simply known as ‘Boneseed’.

C. monilifera was first described by Carl Linnaeus in 1754 under the name Osteospermum monil-
ifera, however it was given its current binomial name in 1943 by T. Norlindh.

The species name monilifera comes from the Latin, monile, meaning necklace or collar, referring
to the shiny fruit arranged around the flowers like a necklace.

Description
Boneseed is a perennial, woody, upright shrub, growing to 3 m (9.8 ft), although occasionally tall-
er. It is a member of the Asteraceae (daisy) family and has showy, bright yellow flowers in swirls of
5-8 ‘petals’ (ray florets) up to 30 mm (1.2 in) in diameter. Fruit are berry-like, spherical at around
8 mm in diameter, and turn dark-brown to black with a bone-coloured seed inside of 6–7 mm di-
ameter. Leaves are 2–6 cm (0.79–2.36 in) long by 1.5–5 cm (0.59–1.97 in) wide, oval tapering to
the base with irregularly serrate margins.

Bitou Bush can be distinguished from Boneseed in part due to its more rounded sprawling habit
to 1.5–2 m (4.9–6.6 ft), less noticeably toothy leaf margins and seeds that are egg-like rather than
spherical.

Both Boneseed and Bitou Bush hybridise readily, however, so examples of plants demonstrating a
fusion of traits is possible.

Distribution and Habitat


Chrysanthemoides monilifera occurs naturally in coastal areas of South Africa, reaching into
southern Namibia and Mozambique. The most widespread subspecies in South Africa is pisifera.
Subspecies rotundata is concentrated along the eastern coast of South Africa from its southern tip

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132 Fundamentals of Weed Science

through to the Mozambique border. Subspecies monilifera is concentrated around Cape Town and
the Cape Peninsula on South Africa’s south western coast.

Invasive Species
Australia
In Australia, subspecies rotundata (Bitou Bush) has naturalised along the coast of Queensland
and New South Wales, while subspecies monilifera (Boneseed) has naturalised along and near the
coast in parts of Victoria and South Australia.

New Zealand
In New Zealand C. monilifera, which is listed on the National Pest Plant Accord, is common in
coastal locations throughout the North Island, and can also be found in the South Island in Nelson
City, Port Hills (Christchurch) and the Otago Peninsula.

Impact
In Australia, C. monilifera has been particularly successful in invading natural bushland. In part,
this is due to the species’ ability to establish on relatively nutrient-poor soils and in areas exposed
to salt such as coastlines, as well as the ability of the seeds to germinate readily. Disturbances such
as fire can assist C. monilifera to spread as the plant produces a large amount of seed that can
persist in the soil seed bank for 10 years or more, and this reserve in turn enables the species to
quickly recolonize a burnt area.

An individual plant can produce 50,000 seeds a year, about 60% of which are viable. Once germinated,
seedlings grow vigorously with dense, bushy growth. This lush growth shades out and displaces slower
growing native species that might otherwise occupy the same ecological niche. Rapid, vigorous growth
also means that C. monilifera is capable of flowering and setting seed within 12–18 months, making it
extremely persistent even in situations where disturbance or regular management activity is common.

Once established, the plant’s shallow root system enables it to absorb moisture after light rain
before the moisture reaches the roots of more deeply rooted species further limiting opportunities
for slower growing species to establish and out-compete C. monilifera over time. Furthermore,
outside of Southern Africa the plant has few local, indigenous pathogens or predators to control its
growth also reducing the potential for gaps to emerge that might provide opportunities for other
species to reestablish. The net consequence of C. monilifera’s growth characteristics is that outside
of its natural ecosystem it can ultimately form large, dense, unhealthy stands of a single species
with extraordinarily poor biodiversity.

The plant can extend its existing range in a variety of ways. Its fruit is attractive to birds, rabbits, other
animals and even some insects such as ants, and because seeds are tough and difficult to digest they
will often be dispersed in animal droppings. Seeds can also spread on vehicles and equipment, in con-
taminated soil, in garden waste, along water drainage lines and deliberately by human intervention.

C. monilifera, unlike many other weed species, is not generally considered to be a problem for
agricultural productivity due to its sensitivity to trampling as well as being readily grazed by stock.

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Species of Weed: Invasive and Introduced 133

Control
C. monilifera is potentially susceptible to a range of control strategies, however Burgman and Lin-
denmayer recommended that the strategy chosen be responsive to the local situation and available
resources. Due to its relatively shallow root system removal by hand is an ideal method of control.
Where manual removal is impractical many common herbicides can be used, in which case the
herbicide is commonly applied directly to the wood of the plant via a cut notch or at the end of
a pruned stump. Mechanical removal of C. monilifera by tractor or other machinery can also be
effective, however this can be extremely indiscriminate and is only recommended in areas of poor
environmental values and minimal erosion risk.

Another method of control available is the use of controlled burns, however there are risks associ-
ated with this method. Principally, C. monilifera has higher moisture levels than many Australian
indigenous species and therefore in Australia a higher than normal intensity fire is required for
burns to be effective. This can in turn have detrimental impacts on indigenous vegetation evolved
in response to more frequent, lower intensity fire. Furthermore, fire can trigger regermination
from the extensive C. monilifera seed bank, potentially worsening the situation unless a program
is implemented to monitor and control C. monilifera seedlings following the burn. If C. monilifera
seedlings are removed, however, this can be extremely effective at exhausting the seed bank and
minimising the chances of reinfestation.

Various methods of biological control have been attempted, particularly the introduction of insects
adapted as natural enemies of C. monilifera such as the Bitou Tip Moth (Comostolopsis germa-
na) and Bitou Seed Fly (Mesoclanis polana). In Australia, while these have had some success in
controlling Bitou Bush (ssp. rotundata), to date they have not had similar success in combating
Boneseed (ssp. monilifera).

Heracleum Mantegazzianum
Heracleum mantegazzianum, commonly known as giant hogweed, cartwheel-flower, giant cow
parsnip, hogsbane or giant cow parsley, is a plant in the family Apiaceae. In New Zealand, it is also
sometimes called wild parsnip, or wild rhubarb. It typically grows to heights of 2–5.5 m (6 ft 7 in–
18 ft 1 in). Superficially, it resembles common hogweed (Heracleum sphondylium), Heracleum
sosnowskyi, or garden angelica (Angelica archangelica). It is phototoxic and considered to be a
noxious weed in many jurisdictions. Giant hogweed is native to the Caucasus region and Central
Asia. It was introduced to Britain as an ornamental plant in the 19th century, and it has also spread
to many other parts of Europe, the United States, and Canada.

The sap of giant hogweed causes phytophotodermatitis in humans, resulting in blisters and
long-lasting scars. These serious reactions are due to the furocoumarin derivatives in the leaves,
roots, stems, flowers, and seeds of the plant.

Description
Giant hogweed has a stout, bright green stem that is frequently spotted with dark red and hollow

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134 Fundamentals of Weed Science

red-spotted leaf stalks that produce sturdy bristles. The stems grow to more than 2 m high. The hol-
low stems vary from 3–8 cm (1.2–3.1 in) in diameter, occasionally up to 10 cm (3.9 in). Each dark
red spot on the stem surrounds a hair, and large, coarse white hairs occur at the base of the leaf stalk.
The plant has deeply incised compound leaves which grow up to 1–1.7 m (3 ft 3 in–5 ft 7 in) in width.

Giant hogweed is a biennial or monocarpic perennial, the plants usually begin dying after they
have set seed. It usually flowers in its second year from late spring to midsummer, with numerous
white flowers clustered in an umbrella-shaped head that is up to 80 cm (31 in) in diameter across
its flat top. The plant produces 1,500 to 100,000 flattened, 1-centimetre (0.39 in)-long, oval, dry
seeds that have a broadly rounded base and broad marginal ridges. Tall dead stems may mark its
locations during winter.

Introduction to Western Europe and North America

Distribution of giant hogweed in Europe (2005)

Giant hogweed was among many foreign plants introduced to Britain in the 19th century as orna-
mental plants. It is now widespread throughout the British Isles, especially along riverbanks. By
forming dense stands, they can displace native plants and reduce wildlife habitats. It has spread in
the northeastern and northwestern United States, and southern Canada and is an invasive species
in Germany, France, and Belgium, overtaking the local native species, Heracleum sphondylium.

In Canada, the plant has been sighted in British Columbia, Alberta, Saskatchewan, Ontario, Que-
bec, New Brunswick, Nova Scotia, and in isolated areas of Newfoundland. It has been seen in
Quebec since the early 1990s. The plant’s spread in Ontario began in the southwest and was seen
in 2010 in the greater Toronto area and Renfrew County near Ottawa.

Giant hogweed was introduced into New York about 1917, and was recorded in British Columbia in
the 1930s. It now occurs in the west in British Columbia, Washington, and Oregon and in eastern
North America from Newfoundland and Nova Scotia west to Ontario and Wisconsin and south to
Indiana, Maryland, and New Jersey. It is also recorded occasionally in Michigan It is a federally
listed noxious weed in many states.

Phototoxicity
The sap of the giant hogweed plant is phototoxic; when the contacted skin is exposed to sunlight
or to ultraviolet rays, it can cause phytophotodermatitis (severe skin inflammations). Initially, the

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Species of Weed: Invasive and Introduced 135

skin colours red and starts itching. Blisters form as it burns within 48 hours. They form black or
purplish scars that can last several years. Hospitalisation may be necessary. Although many media
reports on giant hogweed suggest the plant can lead to temporary or permanent blindness, existing
research on the plant does not back up this claim.

Giant hogweed (close-up)

These reactions are caused by the presence of linear derivatives of furanocoumarin in its leaves,
roots, stems, flowers, and seeds. These chemicals can get into the nucleus of the epithelial cells,
forming a bond with the DNA, causing the cells to die. The brown colour is caused by the produc-
tion of melanin by furocoumarins.

Authorities advise that children should be kept away from giant hogweed, that protective clothing,
including eye protection, should be worn when handling or digging it, and that if skin is exposed,
the affected area should be washed thoroughly with soap and water and the exposed skin protected
from the sun for several days.

Countermeasures
Because of its phototoxicity and invasive nature, giant hogweed is often actively removed. In the
UK, the Wildlife and Countryside Act 1981 makes it an offence to plant or cause giant hogweed to
grow in the wild. Hogweed is regulated as a federal noxious weed by the US government, and is
illegal to import into the United States or move interstate without a permit from the Department of
Agriculture. The USDA Forest Service states pigs and cattle can eat it without apparent harm. The
New York State Department of Environmental Conservation has had an active program to control
giant hogweed since 2008, including reporting, database maintenance, and crews for removal or
herbicide control. In 2011, Maine state horticulturists, describing the plant as “Queen Anne’s lace
on steroids”, reported that it has been found at 21 different locations in Maine, with the number of
plants ranging from one to a hundred.

In Popular Culture
The 1971 album Nursery Cryme by the progressive rock group Genesis contains a song called
“The Return of the Giant Hogweed”. The lyrics describe a murderous attack on the human race by
Heracleum mantegazzianum, long after the plant was first “captured” and brought to England by
a Victorian explorer.

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136 Fundamentals of Weed Science

Impatiens Glandulifera
Impatiens glandulifera is a large annual plant native to the Himalayas. Via human introduction
it is now present across much of the Northern Hemisphere and considered an invasive species in
some areas.

Etymology
The common names Policeman’s Helmet, Bobby Tops, Copper Tops, and Gnome’s Hatstand all
originate from the flowers being decidedly hat-shaped. Himalayan Balsam and Kiss-me-on-the-
mountain arise from the plant originating in the Himalayan mountains. Ornamental jewelweed
refers to its cultivation as an ornamental plant.

The genus name Impatiens, meaning “impatient”, refers to its method of seed dispersal. The spe-
cies name glandulifera comes from the Latin words glandis meaning ‘gland’, and ferre meaning
‘to bear’, referring to the plant’s glands.

Description

Impatiens glandulifera scattering its seeds

Leaves of the Himalayan Balsam

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Species of Weed: Invasive and Introduced 137

It typically grows to 1 to 2 m (3.3 to 6.6 ft) high, with a soft green or red-tinged stem, and lanceolate
leaves 5 to 23 cm (2.0 to 9.1 in) long. The crushed foliage has a strong musty smell. Below the leaf
stems the plant has glands that produce a sticky, sweet-smelling, and edible nectar. The flowers are
pink, with a hooded shape, 3 to 4 cm (1¼ to 1½ in) tall and 2 cm (¾ in) broad; the flower shape
has been compared to a policeman’s helmet.

Glands

After flowering between June and October, the plant forms seed pods 2 to 3 cm (¾ to 1¼ in) long
and 8 mm broad (¼ in), which explode when disturbed, scattering the seeds up to 7 metres (23 feet).

The green seed pods, seeds, young leaves and shoots are all edible. The flowers can be turned into
a jam or parfait.

Distribution
Ireland
Recorded from almost all counties of Ireland.

Invasive Species
Himalayan Balsam is sometimes cultivated for its flowers. It is now widely established in other
parts of the world (such as the British Isles and the United States), in some cases becoming an
invasive species weed. The aggressive seed dispersal, coupled with high nectar production which
attracts pollinators, often allows the Himalayan Balsam to outcompete native plants. Himalayan
Balsam also promotes river bank erosion due to the plant dying back over winter, leaving the bank
unprotected from flooding. Invasive Himalayan Balsam can also adversely affect indigenous spe-
cies by attracting pollinators (e.g. insects) at the expense of indigenous species. It is considered a
“prohibited noxious weed” under the Alberta Weed Control Act 2010.

In the UK the plant was first introduced in 1839 at the same time as Giant Hogweed and Japanese
Knotweed. These plants were all promoted at the time as having the virtues of “herculean propor-
tions” and “splendid invasiveness” which meant that ordinary people could buy them for the cost
of a packet of seeds to rival the expensive orchids grown in the greenhouses of the rich. Within
ten years, however, Himalayan balsam had escaped from the confines of cultivation and begun to
spread along the river systems of England. Today it has spread across most of the UK and some
local wildlife trusts organise “balsam bashing” events to help control the plant. However, a recent

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138 Fundamentals of Weed Science

study (Hejda & Pyšek, 2006) concludes that in some circumstances, such efforts may cause more
harm than good. Destroying riparian stands of Himalayan Balsam can open up the habitat for more
aggressive invasive plants such as Japanese knotweed and aid in seed dispersal (by dropped seeds
sticking to shoes). Riparian habitat is suboptimal for I. glandulifera, and spring or autumn flood-
ing destroys seeds and plants. The research suggests that the optimal way to control the spread of
riparian Himalayan Balsam is to decrease eutrophication, thereby permitting the better-adapted
local vegetation that gets outgrown by the balsam on watercourses with high nutrient load to re-
bound naturally. They caution that these conclusions do probably not hold true for stands of the
plant at forest edges and meadow habitats, where manual destruction is still the best approach.

Himalayan Balsam at Bank Hall, Bretherton, Lancashire, England

The Bionic Control of Invasive Weeds in Wiesbaden, Germany, is trying to establish a self-suffi-
cient project to conserve their local biodiversity by developing several food products made from
the Impatiens flowers. Eventually, if all goes well, this project will have the Himalayan Balsam
financing its own eradication.

In August 2014, CABI released a rust fungus in Berkshire, Cornwall and Middlesex in the United
Kingdom as part of field trials into the biological control of Himalayan Balsam.

Some research also suggests that I. glandulifera may exhibit allelopathy, in which it excretes tox-
ins that negatively affect neighboring plants, thus increasing its competitive advantage. This would
further support the conclusion that pulling or cutting the plant and leaving it to decompose is not
the best method to control this invasive species.

The Royal Horticultural Society and the Centre for Ecology and Hydrology recommend that pull-
ing and cutting is the main method of non-chemical control, and usually the most appropriate.
Natural Resources Wales has used manual methods such as pulling plants and using strimmers to
largely eradicate Himalayan Balsam from reaches of the River Ystwyth.

Fruits of impatiens glandulifera

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Species of Weed: Invasive and Introduced 139

Fallopia Japonica
Fallopia japonica, synonym Reynoutria japonica, commonly known as Japanese knotweed, is a
large, herbaceous perennial plant of the knotweed and buckwheat family Polygonaceae. It is native
to East Asia in Japan, China and Korea. In North America and Europe the species is very successful
and has been classified as an invasive species in several countries. In Australia, it is illegal to have
any of this species growing on one’s property.

Japanese knotweed has hollow stems with distinct raised nodes that give it the appearance of bamboo,
though it is not closely related. While stems may reach a maximum height of 3–4 m (9.8–13.1 ft) each
growing season, it is typical to see much smaller plants in places where they sprout through cracks in
the pavement or are repeatedly cut down. The leaves are broad oval with a truncated base, 7–14 cm
(2.8–5.5 in) long and 5–12 cm (2.0–4.7 in) broad, with an entire margin. The flowers are small, cream
or white, produced in erect racemes 6–15 cm (2.4–5.9 in) long in late summer and early autumn.

Closely related species include giant knotweed (Fallopia sachalinensis, syn. Polygonum sachalin-
ense) and Russian vine (Fallopia baldschuanica, Polygonum baldschuanicum).

Other English names for Japanese knotweed include fleeceflower, Himalayan fleece vine, monkey-
weed, monkey fungus, Hancock’s curse, elephant ears, pea shooters, donkey rhubarb (although
it is not a rhubarb), sally rhubarb, Japanese bamboo, American bamboo, and Mexican bamboo
(though it is not a bamboo). In Chinese medicine, it is known as Huzhang (Chinese: 虎杖; pinyin:
Hǔzhàng), which translates to “tiger stick”. There are also regional names, and it is sometimes
confused with sorrel. In Japanese, the name is itadori (虎杖, イタドリ?).

Old stems remain in place as new growth appears

A hedgerow made up of roses and Japanese knotweed in Caersws, Wales in 2010

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140 Fundamentals of Weed Science

Erect inflorescence

Invasive Species
It is listed by the World Conservation Union as one of the world’s worst invasive species.

The invasive root system and strong growth can damage concrete foundations, buildings, flood
defences, roads, paving, retaining walls and architectural sites. It can also reduce the capacity of
channels in flood defences to carry water.

It is a frequent colonizer of temperate riparian ecosystems, roadsides and waste places. It


forms thick, dense colonies that completely crowd out any other herbaceous species and is
now considered one of the worst invasive exotics in parts of the eastern United States. The
success of the species has been partially attributed to its tolerance of a very wide range of soil
types, pH and salinity. Its rhizomes can survive temperatures of −35 °C (−31 °F) and can ex-
tend 7 metres (23 ft) horizontally and 3 metres (9.8 ft) deep, making removal by excavation
extremely difficult.

The plant is also resilient to cutting, vigorously resprouting from the roots. The most effective
method of control is by herbicide application close to the flowering stage in late summer or au-
tumn. In some cases, it is possible to eradicate Japanese knotweed in one growing season using
only herbicides. Trials in Haida Gwaii, British Columbia, using sea water sprayed on the foliage,
have demonstrated promising results, which may prove to be a viable option for eradication where
concerns over herbicide application are too great.

Two biological pest control agents that show promise in the control of the plant are the psyllid
Aphalara itadori and a leaf spot fungus from genus Mycosphaerella.

New Zealand
Japanese knotweed is classed as an unwanted organism in New Zealand and is established in some
parts of the country.

United Kingdom
European adventurer Philipp Franz von Siebold transported Japanese knotweed from a Japanese
volcano to Holland. By 1850 a specimen from this plant was added to the Royal Botanic Gar-

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Species of Weed: Invasive and Introduced 141

dens, Kew. It was favoured by gardeners because it looked like bamboo and grew everywhere. Ann
Conolly provided the first authoritative work on the history and distribution of the plant in the UK
and Europe in the 1970s. Some home owners in the United Kingdom are unable to sell their homes
if there is any evidence of knotweed on the property.

In the UK, Japanese knotweed is established in the wild in many parts of the country and creates
problems due to the impact on biodiversity, flooding management and damage to property. It is an
offence under section 14(2) of the Wildlife and Countryside Act 1981 to “plant or otherwise cause
to grow in the wild” any plant listed in Schedule nine, Part II to the Act, which includes Japanese
knotweed. It is also classed as “controlled waste” in Britain under part 2 of the Environmental
Protection Act 1990. This requires disposal at licensed landfill sites.

The species is expensive to remove. According to the UK government, the cost of controlling knot-
weed has hit $3 billion. It cost $130 million to eradicate knotweed on a patch of land on the pro-
posed site of the London’s 2012 Olympic Games velodrome and aquatic centre. Defra’s Review of
Non-native Species Policy states that a national eradication programme would be prohibitively
expensive at £1.56 billion.

The decision was taken on 9 March 2010 in the UK to release into the wild a Japanese psyllid in-
sect, Aphalara itadori. Its diet is highly specific to Japanese knotweed and shows good potential
for its control. Controlled release trials began in South Wales in 2016.

In Scotland, the Wildlife and Natural Environment (Scotland) Act 2011 came into force in July
2012 that superseded the Wildlife and Countryside Act 1981. This act states that is an offence to
spread intentionally or unintentionally Japanese knotweed (or other non-native invasive spe-
cies).

United States
The weed can be found in 39 of the 50 United States. It is listed as an invasive weed in Maine, New
Hampshire, Ohio, Vermont, Virginia, West Virginia, New York, New Jersey, Alaska, Pennsylvania,
Michigan, Oregon, Washington state, Wisconsin, Minnesota and Colorado.

Canada
According to Gail Wallin, executive director of the Invasive Species Council of B.C., and co-chair of
the Canadian Council on Invasive Species, by 2015 it was found in all provinces in Canada except
Manitoba and Saskatchewan. In Vancouver the aggressive plant went under “four lanes of high-
way and have popped up on the other side.” At Mission Point Park in Davis Bay, British Columbia
municipal crews attempted to eradicate it by digging out the plant to a depth of about three metres
with an excavator. It grew back twice as large the next year. To avoid an epidemic as in the United
Kingdom, some provinces in Canada are pushing for relaxation of provincial limits on the use of
herbicides close to waterways so knotweed can be aggressively managed with strong chemicals. In
spite of its status as an invasive species it is still sometimes sold or swapped in Canada as an edible
“false bamboo.” Bohemian knotweed, a mutant hybrid— which is a cross between Japanese and
giant knotweed that produces huge quantities of viable seeds —now accounts for about 80 per cent
of knotweed infestations in British Columbia.

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142 Fundamentals of Weed Science

Uses

A variegated variety of Japanese knotweed, used as a landscape plant

Japanese knotweed flowers are valued by some beekeepers as an important source of nectar for
honeybees, at a time of year when little else is flowering. Japanese knotweed yields a monofloral
honey, usually called bamboo honey by northeastern U.S. beekeepers, like a mild-flavored version
of buckwheat honey (a related plant also in the Polygonaceae).

The young stems are edible as a spring vegetable, with a flavor similar to extremely sour rhubarb.
In some locations, semi-cultivating Japanese knotweed for food has been used as a means of con-
trolling knotweed populations that invade sensitive wetland areas and drive out the native vegeta-
tion. It is eaten in Japan as sansai or wild foraged vegetable.

Both Japanese knotweed and giant knotweed are important concentrated sources of resveratrol
and its glucoside piceid. The plant is an available source of resveratrol because of its year-round
growth and robustness in different climates.

This antique locomotive at Beekbergen, Netherlands is overgrown by knotweed. A few years


before, it was free of knotweed

Control
Japanese knotweed has a large underground network of roots (rhizomes). To eradicate the plant
the roots need to be killed. All above-ground portions of the plant need to be controlled repeatedly
for several years in order to weaken and kill the entire patch. Picking the right herbicide is essen-
tial, as it must travel through the plant and into the root system below.

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Species of Weed: Invasive and Introduced 143

The abundance of the plant can be significantly reduced by applying glyphosate, imazapyr, a com-
bination of both, or by cutting all visible stalks and filling the stems with glyphosate. However,
these methods have not been proven to provide reliable long-term results in completely eliminat-
ing the treated population.

Digging up the rhizomes is a common solution where the land is to be developed, as this is quicker
than the use of herbicides, but safe disposal of the plant material without spreading it is difficult;
knotweed is classed as controlled waste in the UK, and disposal is regulated by law. Digging up the
roots is also very labor-intensive and not always efficient. The roots can go to up to 10 feet (3.0 me-
ters) deep, and leaving only a few inches of root behind will result in the plant quickly growing back.

Covering the affected patch of ground with a non-translucent material can be an effective fol-
low-up strategy. However, the trimmed stems of the plant can be razor sharp and are able to pierce
through most materials. Covering with non-flexible materials such as concrete slabs has to be done
meticulously and without leaving even the smallest splits. The slightest opening can be enough for
the plant to grow back.

More ecologically-friendly means are being tested as an alternative to chemical treatments. Soil
steam sterilization involves injecting steam into contaminated soil in order to kill subterranean
plant parts. Research has also been carried out on Mycosphaerella leafspot fungus, which devas-
tates knotweed in its native Japan. This research has been relatively slow due to the complex life
cycle of the fungus.

Research has been carried out by not-for-profit inter-governmental organisation CABI in the UK.
Following earlier studies imported Japanese knotweed psyllid insects (Aphalara itadori), whose
only food source is Japanese knotweed, were released at a number of sites in Britain in a study
running from 1 April 2010 to 31 March 2014. In 2012, results suggested that establishment and
population growth were likely, after the insects overwintered successfully.

Anecdotal reports of effective control describe the use of goats to eat the plant parts above ground
followed by the use of pigs to root out and eat the underground parts of the plant.

Detail of the stalk

Controversy
In the United Kingdom, Japanese knotweed has received a lot of attention in the press as a re-
sult of very restrictive lending policies by banks and other mortgage companies. Several lenders

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144 Fundamentals of Weed Science

have refused mortgage applications on the basis of the plant being discovered in the garden or
neighbouring garden. The Royal Institution of Chartered Surveyors published a report in 2012 in
response to lenders refusing to lend “despite [knotweed] being treatable and rarely causing severe
damage to the property”.

There is a real lack of information and understanding of what Japanese knotweed is and the actual
damage it can cause. Without actual advice and guidance, surveyors have been unsure of how to
assess the risk of Japanese knotweed, which can result in inconsistent reporting of the plant in
mortgage valuations. RICS hopes that this advice will provide the industry with the tools it needs
to measure the risk effectively, and provide banks with the information they require to identify
who and how much to lend to at a time when it is essential to keep the housing market moving.

— Philip Santo, RICS Residential Professional Group

In response to this guidance, several lenders have relaxed their criteria in relation to discovery of
the plant. As recently as 2012, the policy at the Woolwich (part of Barclays plc) was “if Japanese
knotweed is found on or near the property then a case will be declined due to the invasive nature of
the plant.” Their criteria have since been relaxed to a category-based system depending on whether
the plant is discovered on a neighbouring property (categories 1 and 2) or the property itself (cate-
gories 3 and 4) incorporating proximity to the property curtilage and the main buildings. Even in a
worst-case scenario (category 4), where the plant is “within 7 metres of the main building, habitable
spaces, conservatory and/or garage and any permanent outbuilding, either within the curtilage of
the property or on neighbouring land; and/or is causing serious damage to permanent outbuildings,
associated structures, drains, paths, boundary walls and fences” Woolwich lending criteria now spec-
ify that this property may be acceptable if “remedial treatment by a Property Care Association (PCA)
registered firm has been satisfactorily completed. Treatment must be covered by a minimum 10-year
insurance-backed guarantee, which is property specific and transferable to subsequent owners and
any mortgagee in possession.” Santander have relaxed their attitude in a similar fashion.

Property Care Association chief executive Steve Hodgson, whose trade body has set up a task force
to deal with the issue, said: “Japanese knotweed is not ‘house cancer’ and could be dealt with in the
same way qualified contractors dealt with faulty wiring or damp.”

Japan
The plant is known as itadori (虎杖?). The kanji expression is from the Chinese meaning “tiger
staff”. One interpretation of the Japanese name is that it comes from “remove pain” (alluding to its
painkilling use), though there are other etymological explanations offered.

It grows widely throughout Japan and is foraged as a wild edible vegetable (sansai), though not in
sufficient quantities to be included in statistics. They are called by such regional names as tonkiba
(Yamagata), itazuiko (Nagano, Mie), itazura (Gifu, Toyama, Nara, Wakayama, Kagawa), gonpachi
(Shizuoka, Nara, Mie, Wakayama), sashi (Akita, Yamagata), jajappo (Shimane, Tottori, Okaya-
ma), sukanpo (many areas).

Young leaves and shoots, which look like asparagus, are used. They are extremely sour; the fibrous
outer skin must be peeled, soaked in water for half a day raw or after parboiling, before being
cooked.

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Species of Weed: Invasive and Introduced 145

Places in Shikoku such as central parts of Kagawa Prefecture pickle the peeled young shoots by
weighting them down in salt mixed with 10% nigari (magnesium chloride). People in Kochi also
rub these cleaned shoots with coarse salt-nigari blend. It is said (though no authority is cited) that
the magnesium of the nigari binds with the oxalic acid thus mitigating its hazard.

Pilosella Aurantiaca
Pilosella aurantiaca (fox-and-cubs, orange hawkweed, tawny hawkweed, devil’s paintbrush, grim-
the-collier) is a perennial flowering plant in the daisy family Asteraceae native to alpine regions of
central and southern Europe, where it is protected in several regions.

Description
It is a low-growing plant with shallow fibrous roots and a basal rosette of elliptical to lanceolate
leaves 5–20 cm long and 1–3 cm broad. All parts of the plant exude a milky juice. The flowering
stem is usually leafless or with just one or two small leaves. The stem and leaves are covered with
short stiff hairs (trichomes), usually blackish in color. The stems may reach a height of 60 cm and
have 2–25 capitula (flowerheads), each 1–2½ cm diameter, bundled together at the end of short
pedicels. The flowers are orange, almost red, which is virtually invisible to bees, yet they also re-
flect ultraviolet light, increasing their conspicuousness to pollinators. The flowers are visited by
various insects, specifically Musca flies. The flowers themselves come in a range of colors from a
deep rust-orange to a pure yellow and often show striking gradients of color.

The plant propagates through its wind-dispersed seeds, and also vegetatively by stolons and shal-
low rhizomes.

Cultivation and Uses

Whole plant

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146 Fundamentals of Weed Science

P. aurantiaca is widely grown as an ornamental plant in gardens for its very decorative flowers. It
has been introduced into Australasia and North America and escaped from gardens and is consid-
ered an invasive species in some areas. It is found across Canada and the north of the U.S., reach-
ing more in the south on the coasts. The plant is easily removed (until it seeds back) by ploughing
or salting, and it cannot be controlled by mowing because it regularly reproduces vegetatively. It is
on the noxious weeds and/or quarantine lists of Colorado, Idaho, Montana, Oregon, Washington,
and regions of British Columbia, with cultivation usually prohibited. In Alberta, it is not consid-
ered a noxious weed except locally. In Australia, the plant is considered a noxious weed in Tasma-
nia and it is quarantined from the country. It has been recorded at Kosciuszko National Park in
New South Wales, Falls Creek and Mount Buller in Victoria and Fern Tree and Snug in Tasmania.
In New Zealand, the plant is closely watched and attempts at control are made.

Echium Plantagineum
Echium plantagineum, commonly known as purple viper’s-bugloss or Paterson’s curse, is a spe-
cies of Echium native to western and southern Europe (from southern England south to Iberia
and east to the Crimea), northern Africa, and southwestern Asia (east to Georgia). It has also been
introduced to Australia, South Africa and United States, where it is an invasive weed. Due to a high
concentration of pyrrolizidine alkaloids, it is poisonous to grazing livestock, especially those with
simple digestive systems, like horses. The toxins accumulate in the liver, and death results from
too much Paterson’s curse in the diet.

Description
Echium plantagineum is a winter annual plant growing to 20–60 cm tall, with rough, hairy, lan-
ceolate leaves up to 14 cm long. The flowers are purple, 15–20 mm long, with all the stamens pro-
truding, and borne on a branched spike.

Invasive Species

In Adelaide, South Australia

Echnium plantagineum has become an invasive species in Australia, where it is also known as
Salvation Jane (particularly in South Australia), blueweed, Lady Campbell weed, and Riverina
bluebell.

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Species of Weed: Invasive and Introduced 147

In the United States the species has become naturalised in parts of California, Oregon, and some
eastern states and areas such as northern Michigan. In Oregon it has been declared a noxious
weed.

Medical Research
In a study funded by the National Center for Complementary and Integrative Health, the seeds
were found to lower triglycerides. Researchers at Wake Forest University and the Harvard Center
for Botanical Lipids fed mice a diet supplemented with echium oil and found that it had effects
similar to fish oil in lowering triglyceride levels in blood plasma and the liver.

Cosmetics
Echium oil contains high levels of alpha linolenic acid (ALA), gamma linolenic acid (GLA), and
stearidonic acid (SDA), making it valuable in cosmetic and skin care applications, with further
potential as an alternative to dietary fish oils.

Toxicity
Echium plantagineum contains pyrrolizidine alkaloids and is poisonous. When eaten in large
quantities, it causes reduced livestock weight and death, in severe cases. Paterson’s curse can
kill horses and irritate the udders of dairy cows and the skin of humans. After the 2003 Can-
berra bushfires a large bloom of the plant occurred on the burned land, and many horses
became ill and died from grazing on it. Because the alkaloids can also be found in the nectar
of Paterson’s curse, the honey made from it should be blended with other honeys to dilute the
toxins.

Asparagus Asparagoides
Asparagus asparagoides, commonly known as Bridal creeper, Bridal-veil creeper, Gnarboola,
Smilax or Smilax asparagus, is a herbaceous climbing plant of the family Asparagaceae native to
eastern and southern Africa. Sometimes grown as an ornamental plant, it has become a serious
environmental weed in Australia and New Zealand.

Taxonomy
Linnaeus first described this species as Medeola asparagoides in 1753. It has been reclassified in
the genus Asparagus by W. Wight in 1909, or Myrsiphyllum by Carl Ludwig von Willdenow in
1808.

Description
Asparagus asparagoides grows as a herbaceous vine with a scrambling or climbing habit which
can reach 3 m (10 ft) in length. It has shiny green leaf-like structures (phylloclades) which are
flattened stems rather than true leaves. They measure up to 4 cm long by 2 cm wide. The pendent

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148 Fundamentals of Weed Science

white flowers appear over winter and spring, from July to September. It is rhizomatous, and bears
tubers which reach 6 cm (2.4 in) by 2 cm (1.8 in) in size.

Distribution and Habitat


It ranges throughout tropical Africa, south to Namibia, and the fynbos in South Africa, as far south
as Cape Town.

It has become naturalised in parts of southern California.

Uses
Asparagus asparagoides, often under the name smilax, is commonly used in floral arrangements
or home decorating.

Invasive Species

A bridal creeper infestation in Australia

A. asparagoides is a major weed species in southern Australia and in New Zealand.

It was introduced to Australia from South Africa around 1857, for use as a foliage plant, especial-
ly in bridal bouquets (hence the common name). It has escaped into the bush and smothers the
native vegetation with the thick foliage and thick underground mat of tubers which restrict root
growth of other species. It is recognised as one of the 20 “weeds of national significance”. The seeds
are readily spread in the droppings of birds, rabbits and foxes, as well as the plant extending its
root system. CSIRO have introduced several biological controls in an attempt to reduce the spread
and impact of the weed.

In New Zealand A. asparagoides is listed under the National Pest Plant Accord and is classified as
an “unwanted organism”.

A. asparagoides has also escaped cultivation in California.

References
• Blood, K (2001), Environmental weeds: a field guide for SE Australia, Melbourne, Vic., Australia: CH Jerram &
Associates, pp. 46–47, 86, ISBN 0-9579086-0-1, OCLC 156877920

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Species of Weed: Invasive and Introduced 149

• Parsons, WT (1973), Noxious weeds of Victoria, Melbourne, Vic., Australia: Inkata Press, pp. 100–101, ISBN
0-909605-00-9, OCLC 874633

• Emert, S (2001), Gardener’s companion to weeds (2nd ed.), Sydney, NSW, Australia: Reed New Holland, p.
100, ISBN 1-876334-77-0, OCLC 52245716

• Stace, C.A. (2010). New flora of the British isles (Third ed.). Cambridge, U.K.: Cambridge University Press. p.
450. ISBN 9780521707725.

• Hackney, P. (Ed)1992. Stewart & Corry’s Flora of the North-East of Ireland. The Institute of Irish Studies, The
Queen’s University of Belfast. ISBN 0 85389 446 9

• Peterson, Roger Tory; McKenny, Margaret (1968). A Field Guide to Wildflowers of Northeastern and North-Cen-
tral North America. Houghton Mifflin Company Boston. ISBN 0-395-18325-1.

• Department of the Environment and Heritage and the CRC for Australian Weed Management (2003). Orange
hawkweed (Hieracium aurantiacum). ISBN 1-920932-25-9.

• Eggli, Urs; Hartmann, Heidrun E. K.; Albers, Focke; Meve, Ulrich. Illustrated handbook of succulent plants. 6.
Springer. p. 231. ISBN 3-540-41692-7.

• Clarke, Oscar F. (2007). Flora of the Santa Ana River and environs: with references to world botany. Heyday.
p. 92. ISBN 1-59714-050-3.

• Van Der Kooi, C. J.; Pen, I.; Staal, M.; Stavenga, D. G.; Elzenga, J. T. M. (2015). “Competition for pollinators
and intra-communal spectral dissimilarity of flowers” (PDF). Plant Biology. doi:10.1111/plb.12328.

• “BSBI List 2007”. Botanical Society of Britain and Ireland. Archived from the original (xls) on 2015-02-25.
Retrieved 2014-10-17.

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4
Plants Considered to be Weed
The plants considered to be weed are amaranth, Toxicodendron radicans, tataxacum, Silybum
marianum and striga. Amaranth is a short-lived perennial plant; most of the species of amaran-
thus are summer annual weeds. The topics discussed in the section are of great importance to
broaden the existing knowledge on weed.

Amaranth
Amaranthus, collectively known as amaranth, is a cosmopolitan genus of annual or short-lived pe-
rennial plants. Some amaranth species are cultivated as leaf vegetables, pseudocereals, and orna-
mental plants. Most of the species from Amaranthus are summer annual weeds and are commonly
referred to as pigweed. Catkin-like cymes of densely packed flowers grow in summer or autumn.
Approximately 60 species are recognized, with inflorescences and foliage ranging from purple and
red to green or gold. Members of this genus share many characteristics and uses with members of
the closely related genus Celosia.

“Amaranth” derives from Greek (amárantos), “unfading,” with the Greek word for “flower,” , fac-
toring into the word’s development as amaranth. The more accurate amarant is an archaic vari-
ant.

Taxonomy

Skull shapes made of amaranth and honey for Day of the Dead in Mexico

Amaranthus shows a wide variety of morphological diversity among and even within certain spe-
cies. Although the family (Amaranthaceae) is distinctive, the genus has few distinguishing char-
acters among the 70 species included. This complicates taxonomy and Amaranthus has generally
been considered among systematists as a “difficult” genus.

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Plants Considered to be Weed 151

Formerly, Sauer (1955) classified the genus into two subgenera, differentiating only between mon-
oecious and dioecious species: Acnida (L.) Aellen ex K.R. Robertson and Amaranthus. Although
this classification was widely accepted, further infrageneric classification was (and still is) needed
to differentiate this widely diverse group.

Traditional Mexican candy made with amaranth

Currently, Amaranthus includes three recognized subgenera and 70 species, although species
numbers are questionable due to hybridization and species concepts. Infrageneric classification
focuses on inflorescence, flower characters and whether a species is monoecious/dioecious, as in
the Sauer (1955) suggested classification. A modified infrageneric classification of Amaranthus
was published by Mosyakin & Robertson (1996) and includes three subgenera: Acnida, Amaran-
thus, and Albersia. The taxonomy is further differentiated by sections within each of the subgen-
era.

Species
Species include:

• Amaranthus acanthochiton – greenstripe

• Amaranthus acutilobus – is a synonym of Amaranthus viridis

• Amaranthus albus – white pigweed, tumble pigweed

• Amaranthus anderssonii

• Amaranthus arenicola – sandhill amaranth

• Amaranthus australis – southern amaranth

• Amaranthus bigelovii – Bigelow’s amaranth

• Amaranthus blitoides – mat amaranth, prostrate amaranth, prostrate pigweed

• Amaranthus blitum – purple amaranth

• Amaranthus brownii – Brown’s amaranth

• Amaranthus californicus – California amaranth, California pigweed

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152 Fundamentals of Weed Science

• Amaranthus cannabinus – tidal-marsh amaranth

• Amaranthus caudatus – love-lies-bleeding, pendant amaranth, tassel flower, quilete

• Amaranthus chihuahuensis – Chihuahuan amaranth

• Amaranthus crassipes – spreading amaranth

• Amaranthus crispus – crispleaf amaranth

• Amaranthus cruentus – purple amaranth, red amaranth, Mexican grain amaranth

• Amaranthus deflexus – large-fruit amaranth

• Amaranthus dubius – spleen amaranth, khada sag

• Amaranthus fimbriatus – fringed amaranth, fringed pigweed

• Amaranthus floridanus – Florida amaranth

• Amaranthus furcatus

• Amaranthus graecizans

• Amaranthus grandiflorus

• Amaranthus greggii – Gregg’s amaranth

• Amaranthus hybridus – smooth amaranth, smooth pigweed, red amaranth

• Amaranthus hypochondriacus – Prince-of-Wales feather, prince’s feather

• Amaranthus interruptus – Australian amaranth

• Amaranthus minimus

• Amaranthus mitchellii

• Amaranthus muricatus – African amaranth

• Amaranthus obcordatus – Trans-Pecos amaranth

• Amaranthus palmeri – Palmer’s amaranth, Palmer pigweed, careless weed

• Amaranthus polygonoides – tropical amaranth

• Amaranthus powellii – green amaranth, Powell amaranth, Powell pigweed

• Amaranthus pringlei – Pringle’s amaranth

• Amaranthus pumilus – seaside amaranth

• Amaranthus retroflexus – red-root amaranth, redroot pigweed, common amaranth

• Amaranthus scleranthoides – variously Amaranthus sclerantoides

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Plants Considered to be Weed 153

• Amaranthus scleropoides – bone-bract amaranth

• Amaranthus spinosus – spiny amaranth, prickly amaranth, thorny amaranth

• Amaranthus standleyanus

• Amaranthus thunbergii – Thunberg’s amaranth

• Amaranthus torreyi – Torrey’s amaranth

• Amaranthus tricolor – Joseph’s-coat

• Amaranthus tuberculatus – rough-fruit amaranth, tall waterhemp

• Amaranthus viridis – slender amaranth, green amaranth

• Amaranthus watsonii – Watson’s amaranth

• Amaranthus wrightii – Wright’s amaranth

Nutrition
1 cup (2.4dl, 245g) of cooked amaranth grain (from approx. 65g raw) provides 251 calories and is
an excellent source (20% or more of the Daily Value, DV) of protein, dietary fiber, and some dietary
minerals. Amaranth is particularly rich in manganese (105% DV), magnesium (40% DV), iron
(29% DV), and selenium (20% DV).

Cooked amaranth leaves are an excellent source of vitamin A, vitamin C, calcium, manganese and
folate.

Amaranth does not contain gluten, so it may be a healthy and less expensive alternative to ingredi-
ents traditionally used in gluten-free products. Amaranth has high biological value and its benefits
are not limited to people with gluten-related disorders, but are applicable to the general popula-
tion. Quantity and quality of proteins of amaranth are superior to that of wheat. It also contains
higher concentrations of folic acid with respect to wheat (102 µg/100 g in amaranth vs. 40 µg/100
g in wheat), and its fiber and minerals content are higher to those of other cereals.

Amaranth contains phytochemicals that may be anti-nutrient factors, such as polyphenols, sapo-
nins, tannins and oxalates which are reduced in content and effect by cooking.

Human Uses
History
Known to the Aztecs as huauhtli, it is thought to have represented up to 80% of their caloric con-
sumption before the Spanish conquest. Another important use of amaranth throughout Meso-
america was to prepare ritual drinks and foods. To this day, amaranth grains are toasted much
like popcorn and mixed with honey, molasses or chocolate to make a treat called alegría, mean-
ing “joy” in Spanish. Diego Durán described the festivities for Huitzilopochtli, the name of which
means “hummingbird of the left side” or “left-handed hummingbird” (Real hummingbirds feed on

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154 Fundamentals of Weed Science

amaranth flowers). The Aztec month of Panquetzaliztli (7 December to 26 December) was dedicat-
ed to Huitzilopochtli. People decorated their homes and trees with paper flags; there were ritual
races, processions, dances, songs, prayers, and finally human sacrifices. This was one of the more
important Aztec festivals, and the people prepared for the whole month. They fasted or ate very
little; a statue of the god was made out of amaranth seeds and honey, and at the end of the month,
it was cut into small pieces so everybody could eat a little piece of the god. After the Spanish con-
quest, cultivation of amaranth was outlawed, while some of the festivities were subsumed into the
Christmas celebration.

Because of its importance as a symbol of indigenous culture, its gluten-free palatability, ease of
cooking, and a protein that is particularly well-suited to human nutritional needs, interest in grain
amaranth (especially A. cruentus and A. hypochondriacus) revived in the 1970s. It was recovered
in Mexico from wild varieties and is now commercially cultivated. It is a popular snack sold in
Mexico, sometimes mixed with chocolate or puffed rice, and its use has spread to Europe and parts
of North America. Amaranth and quinoa are non-grasses and are called pseudocereals because of
their similarities to cereals in flavor and cooking.

Seed
Several species are raised for amaranth “grain” in Asia and the Americas.

Ancient amaranth grains still used to this day include the three species, Amaranthus caudatus,
Amaranthus cruentus, and Amaranthus hypochondriacus. Although amaranth was cultivated on
a large scale in ancient Mexico, Guatemala, and Peru, nowadays it is only cultivated on a small
scale there, along with India, China, Nepal, and other tropical countries; thus, there is potential
for further cultivation in those countries, amaranth was described as “the crop of the future.” It
has been proposed as an inexpensive native crop that could be cultivated by indigenous people in
rural areas for several reasons:

• It is easily harvested.

• Its raw seeds are a good source of protein.

• In cooked and edible forms, amaranth retains adequate content of several dietary minerals.

• It is easy to cook.

• As befits its weedy life history, amaranth grains grow rapidly and, in three cultivated spe-
cies of amaranth, their large seedheads can weigh up to 1 kilogram and contain a half-mil-
lion small seeds.

Seed Flour
Amaranth seed flour has been evaluated as an additive to wheat flour by food specialists. To de-
termine palatability, different levels of amaranth grain flour were mixed with the wheat flour and
baking ingredients (1% salt, 2.5% fat, 1.5% yeast, 10% sugar and 52–74% water), fermented, mold-
ed, pan-proofed and baked. The baked products were evaluated for loaf volume, moisture content,
color, odor, taste and texture. The amaranth containing products were then compared with bread

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Plants Considered to be Weed 155

made from 100% wheat flour. The loaf volume decreased by 40% and the moisture content in-
creased from 22 to 42% with increase in amaranth grain flour. The study found that the sensory
scores of the taste, odor, color, and texture decreased with increasing amounts of amaranth. Gen-
erally, above 15% amaranth grain flour, there were significant differences in the evaluated sensory
qualities and the high amaranth-containing product was found to be of unacceptable palatability
to the population sample that evaluated the baked products.

Leaves, Roots, and Stems


Amaranth species are cultivated and consumed as a leaf vegetable in many parts of the world. Four
species of Amaranthus are documented as cultivated vegetables in eastern Asia: Amaranthus cru-
entus, Amaranthus blitum, Amaranthus dubius, and Amaranthus tricolor.

In Indonesia and Malaysia, leaf amaranth is called bayam. In the Philippines, the Ilocano word for
the plant is “kalunay”; the Tagalog word for the plant is kilitis or kulitis. In the state of Uttar Pradesh
and Bihar in India, it is called chaulai and is a popular green leafy vegetable (referred to in the class
of vegetable preparations called saag). It is called chua in Kumaun area of Uttarakhand, where it is a
popular red-green vegetable. In Karnataka state in India, it is called harive. It is used to prepare cur-
ries like hulee, palya, majjigay-hulee, and so on. In the state of Kerala, it is called cheera and is con-
sumed by stir-frying the leaves with spices and red chillies to make cheera thoran. In Tamil Nadu,
it is called mulaikkira and is regularly consumed as a favourite dish, where the greens are steamed,
and mashed, with light seasoning of salt, red chili, and cumin. It is called keerai masial. In Andhra
Pradesh, this leaf is added in preparation of a popular dal called thotakura pappu in (Telugu). In
Maharashtra, it is called shravani maath and is available in both red and white colour. In Orissa, it
is called khada saga, it is used to prepare saga bhaja, in which the leaf is fried with chili and onions.

In China, the leaves and stems are used as a stir-fry vegetable, or in soups. In Vietnam, it is called
rau dền and is used to make soup. Two species are popular as edible vegetable in Vietnam: dền
đỏ- Amaranthus tricolor and dền cơm or dền trắng- Amaranthus viridis.

A traditional food plant in Africa, amaranth has the potential to improve nutrition, boost food se-
curity, foster rural development and support sustainable land care.

In Bantu regions of Uganda and Western Kenya, it is known as doodo or litoto. It is also known
among the Kalenjin as a drought crop (chepkerta). In Nigeria, it is a common vegetable and goes
with all Nigerian starch dishes. It is known in Yoruba as shoko a short form of shokoyokoto
(meaning make the husband fat) or arowo jeja (meaning “we have money left over for fish”). In
the Caribbean, the leaves are called bhaji in Trinidad and callaloo in Jamaica, and are sautéed
with onions, garlic, and toma-toes, or sometimes used in a soup called pepperpot soup. In
Botswana, it is referred to as morug and cooked as a staple green vegetable.

In Greece, green amaranth (A. viridis) is a popular dish called vlita or vleeta. It is boiled, then
served with olive oil and lemon juice like a salad, sometimes alongside fried fish. Greeks stop
harvesting the plant (which also grows wild) when it starts to bloom at the end of August.

In Brazil, green amaranth was, and to a degree still is, frequently regarded as an invasive species
as all other species of amaranth (except the generally imported A. caudatus cultivar), though some

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156 Fundamentals of Weed Science

have traditionally appreciated it as a leaf vegetable, under the names of caruru or bredo, which is
consumed cooked, generally accompanying the staple food, rice and beans.

Dyes
The flowers of the ‘Hopi Red Dye’ amaranth were used by the Hopi (a tribe in the western United
States) as the source of a deep red dye. Also a synthetic dye was named “amaranth” for its similar-
ity in color to the natural amaranth pigments known as betalains. This synthetic dye is also known
as Red No. 2 in North America and E123 in the European Union.

Ornamentals

Amaranthus flowering

The genus also contains several well-known ornamental plants, such as Amaranthus caudatus
(love-lies-bleeding), a vigorous, hardy annual with dark purplish flowers crowded in handsome
drooping spikes. Another Indian annual, A. hypochondriacus (prince’s feather), has deeply veined
lance-shaped leaves, purple on the under face, and deep crimson flowers densely packed on erect
spikes.

Amaranths are recorded as food plants for some Lepidoptera (butterfly and moth) species includ-
ing the nutmeg moth and various case-bearer moths of the genus Coleophora: C. amaranthella, C.
enchorda (feeds exclusively on Amaranthus), C. immortalis (feeds exclusively on Amaranthus),
C. lineapulvella and C. versurella (recorded on A. spinosus).

Ecology
Amaranth weed species have an extended period of germination, rapid growth, and high rates of
seed production, and have been causing problems for farmers since the mid-1990s. This is par-
tially due to the reduction in tillage, reduction in herbicidal use and the evolution of herbicidal
resistance in several species where herbicides have been applied more often. The following 9 spe-
cies of Amaranthus are considered invasive and noxious weeds in the U.S and Canada: A. albus,
A. blitoides, A. hybridus, A. palmeri, A. powellii, A. retroflexus, A. spinosus, A. tuberculatus, and
A. viridis.

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Plants Considered to be Weed 157

A new herbicide-resistant strain of Amaranthus palmeri has appeared; it is glyphosate-resistant


and so cannot be killed by herbicides using the chemical. Also, this plant can survive in tough con-
ditions.This could be of particular concern to cotton farmers using glyphosate-resistant cotton.
The species Amaranthus palmeri (Palmer amaranth) causes the greatest reduction in soybean
yields and has the potential to reduce yields by 17-68% in field experiments. Palmer amaranth is
among the “top five most troublesome weeds” in the southeast of the United States and has already
evolved resistances to dinitroaniline herbicides and acetolactate synthase inhibitors. This makes
the proper identification of Amaranthus species at the seedling stage essential for agriculturalists.
Proper weed control needs to be applied before the species successfully colonizes in the crop field
and causes significant yield reductions.

Myth, Legend and Poetry


The word amaranth comes from the Greek word amaranton, meaning “unwilting” (from the verb
marainesthai meaning “wilt”). The word was applied to amaranth because it did not soon fade
and so symbolized immortality. “Amarant” is a more correct, albeit archaic form, chiefly used in
poetry. The current spelling, amaranth, seems to have come from folk etymology that assumed the
final syllable derived from the Greek word anthos (“flower”), common in botanical names.

An early Greek fable counted among Aesop’s Fables compares the rose to the amaranth to illus-
trate the difference in fleeting and everlasting beauty:

An amaranth planted in a garden near a Rose-Tree, thus addressed it: “What a lovely flower is the
Rose, a favourite alike with Gods and with men. I envy you your beauty and your perfume.” The
Rose replied, “I indeed, dear Amaranth, flourish but for a brief season! If no cruel hand pluck me
from my stem, yet I must perish by an early doom. But thou art immortal and dost never fade, but
bloomest for ever in renewed youth.”

John Milton’s epic Paradise Lost gives amaranth an illustrious neighbour:

Immortal amaranth, a flower which once In paradise, fast by the tree of life, Began to bloom; but
soon for man’s offence To heaven removed, where first it grew, there grows, And flowers aloft,
shading the fount of life, And where the river of bliss through midst of heaven Rolls o’er elysian
flowers her amber stream: With these that never fade the spirits elect Bind their resplendent locks.

Samuel Taylor Coleridge, in Work Without Hope (1825), also refers to the herb, likely referencing
Milton’s earlier work. (ll 7-10 excerpted):

Yet well I ken the banks where Amaranths blow, Have traced the fount whence streams of nectar flow.
Bloom, O ye Amaranths! bloom for whom ye may, For me ye bloom not! Glide, rich streams, away!

Percy Bysshe Shelley refers to the herb in his poem “Bereavement” (Lines 13-16 excerpted):

Eternity points, in its amaranth bower Where no clouds of fate o’er the sweet prospect lour, Un-
speakable pleasure, of goodness the dower, When woe fades away like the mist of the heath.

In his dialogue “Aesop and Rhodopè”, published in 1844, Walter Savage Landor wrote:

There are no fields of amaranth on this side of the grave: there are no voices, O Rhodopè, that are

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158 Fundamentals of Weed Science

not soon mute, however tuneful: there is no name, with whatever emphasis of passionate love re-
peated, of which the echo is not faint at last.

Joachim du Bellay mentioned the herb in his “A Vow To Heavenly Venus,” ca. 1500:

We that with like hearts love, we lovers twain, New wedded in the village by thy fane, Lady of all
chaste love, to thee it is We bring these amaranths, these white lilies, A sign, and sacrifice; may
Love, we pray, Like amaranthine flowers, feel no decay; Like these cool lilies may our loves remain,
Perfect and pure, and know not any stain; And be our hearts, from this thy holy hour, Bound each
to each, like flower to wedded flower.

In the fourth book of Endymion (1818), John Keats writes:

The spirit culls Unfaded amaranth, when wild it strays Through the old garden-ground of boyish
days.

In ancient Greece, the amaranth (also called chrysanthemum and helichrysum) was sacred to
Ephesian Artemis. It allegedly had special healing properties, and, as a symbol of immortality,
was used to decorate images of the gods and tombs. In legend, Amarynthus (a form of “Amaran-
tus”) was a hunter of Artemis and king of Euboea; in a village of Amarynthus, of which he was the
eponymous hero, there was a famous temple of Artemis Amarynthia or Amarysia (Strabo x. 448;
Pausan. i. 31, p. 5). The Chinese used amaranth widely for its healing chemicals, treating illnesses
such as infections, rashes, and migraines. “Amarantos” is the name of a several-centuries-old pop-
ular Greek folk-song:

Look at the amaranth: on tall mountains it grows, on the very stones and rocks and places inac-
cessible.

In the poem “The Hound of Heaven” (1893), Francis Thompson compares God’s love to “an ama-
ranthine weed...”

Ah! is Thy love indeed A weed, albeit an amaranthine weed, Suffering no flowers except its own to
mount?

The Swedish metal band Amaranthe is named for the plant.

Images

Love-lies-bleeding (Amaranthus caudatus)

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Plants Considered to be Weed 159

Green Amaranth (A. hybridus)

Seabeach amaranth (A. pumilus), an amaranth on the Federal Threatened species List

Red-root Amaranth (A. retroflexus) - from Thomé, Flora von Deutschland, Österreich und der Schweiz 1885

Toxicodendron Radicans
Toxicodendron radicans, commonly known as eastern poison ivy or poison ivy, is a poisonous
Asian and North American flowering plant that is well known for causing an itching, irritating, and
sometimes painful rash in most people who touch it, caused by urushiol, a clear liquid compound
in the plant’s sap. The species is variable in its appearance and habit, and despite its common
name it is not a true ivy (Hedera), but rather a member of the cashew family. Toxicodendron rad-
icans is commonly eaten by many animals, and the seeds are consumed by birds, but poison ivy is
most often thought of as an unwelcome weed.

Description
There are numerous subspecies and/or varieties of T. radicans, which can be found growing in any
of the following forms; all of which have woody stems:

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160 Fundamentals of Weed Science

• as a climbing vine that grows on trees or some other support

• as a shrub up to 1.2 metres (3 ft 11 in) tall

• as a trailing vine that is 10–25 centimetres (3.9–9.8 in) tall

Subspecies and varieties

• Toxicodendron radicans subsp. eximum (Greene) Gillis

• Toxicodendron radicans subsp. hispidum (Engl.) Gillis

• Toxicodendron radicans subsp. negundo (Greene) Gillis

• Toxicodendron radicans var. negundo (Greene) Reveal

• Toxicodendron radicans var. pubens (Engelm. ex S. Watson) Reveal

• Toxicodendron radicans subsp. radicans

• Toxicodendron radicans var. radicans

• Toxicodendron radicans subsp. rydbergii (Small ex Rydb.) Á. Löve & D. Löve

• Toxicodendron radicans var. rydbergii (Small ex Rydb.) Erskine

• Toxicodendron radicans subsp. verrucosum (Scheele) Gillis

The deciduous leaves of T. radicans are trifoliate with three almond-shaped leaflets. Leaf color
ranges from light green (usually the younger leaves) to dark green (mature leaves), turning bright
red in fall; though other sources say leaves are reddish when expanding, turn green through ma-
turity, then back to red, orange, or yellow in the fall. The leaflets of mature leaves are somewhat
shiny. The leaflets are 3–12 cm (1.2–4.7 in) long, rarely up to 30 cm (12 in). Each leaflet has a few
or no teeth along its edge, and the leaf surface is smooth. Leaflet clusters are alternate on the vine,
and the plant has no thorns. Vines growing on the trunk of a tree become firmly attached through
numerous aerial rootlets. The vines develop adventitious roots, or the plant can spread from rhi-
zomes or root crowns. The milky sap of poison ivy darkens after exposure to the air.

The urushiol compound in poison ivy is not a defensive measure; rather, it helps the plant to retain
water. It is frequently eaten by animals such as deer and bears.

Toxicodendron radicans vine with typical reddish “hairs.” Like the leaves, the vines are poisonous to humans.

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Plants Considered to be Weed 161

Toxicodendron radicans spreads either vegetatively or sexually. It is dioecious; flowering occurs


from May to July. The yellowish- or greenish-white flowers are typically inconspicuous and are
located in clusters up to 8 cm (3.1 in) above the leaves. The berry-like fruit, a drupe, mature by
August to November with a grayish-white colour. Fruits are a favorite winter food of some birds
and other animals. Seeds are spread mainly by animals and remain viable after passing through
the digestive tract.

Toxicodendron radicans in Perrot State Park, Trempealeau County, Wisconsin

Flower detail, with bee

Distribution and Habitat


Toxicodendron radicans grows throughout much of North America, including the Canadian Mar-
itime provinces, Quebec, Ontario, Manitoba, and all U.S. states east of the Rocky Mountains, as
well as in the mountainous areas of Mexico up to around 1,500 m (4,900 ft). Caquistle or caxuistle
is the Nahuatl term for the species. It is normally found in wooded areas, especially along edge ar-
eas where the tree line breaks and allows sunshine to filter through. It also grows in exposed rocky
areas, open fields and disturbed areas.

Eastern Poison Ivy (Toxicodendron radicans) range map.

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162 Fundamentals of Weed Science

It may grow as a forest understory plant, although it is only somewhat shade-tolerant. The plant
is extremely common in suburban and exurban areas of New England, the Mid-Atlantic, and the
Southeastern United States. The similar species T. diversilobum (western poison oak) and T. ryd-
bergii (western poison ivy) are found in western North America.

Toxicodendron radicans rarely grows at altitudes above 1,500 m (4,900 ft), although the altitude
limit varies in different locations. The plants can grow as a shrub up to about 1.2 metres (3.9 ft)
tall, as a groundcover 10–25 cm (3.9–9.8 in) high, or as a climbing vine on various supports. Older
vines on substantial supports send out lateral branches that may be mistaken for tree limbs at first
glance.

It grows in a wide variety of soil types, and soil pH from 6.0 (acidic) to 7.9 (moderately alkaline). It
is not particularly sensitive to soil moisture, although it does not grow in desert or arid conditions.
It can grow in areas subject to seasonal flooding or brackish water.

It is more common now than when Europeans first arrived in North America. The development of
real estate adjacent to wild, undeveloped land has engendered “edge effects”, enabling poison ivy
to form vast, lush colonies in these areas. It is listed as a noxious weed in the US states of Minne-
sota and Michigan and in the Canadian province of Ontario.

Outside North America, T. radicans is also found in the temperate parts of Asia, in Japan, Taiwan,
the Russian islands of Sakhalin and the Kuriles, and in parts of China.

A study by researchers at the University of Georgia found that poison ivy is particularly sensitive
to carbon dioxide levels, greatly benefiting from higher concentrations in the atmosphere. Poison
ivy’s growth and potency has already doubled since the 1960s, and it could double again once car-
bon dioxide levels reach 560 ppm.

Aids to Identification

T. radicans leaf and berries

T. radicans foliage

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Plants Considered to be Weed 163

The following four characteristics are sufficient to identify poison ivy in most situations: (a) clus-
ters of three leaflets, (b) alternate leaf arrangement, (c) lack of thorns, and (d) each group of three
leaflets grows on its own stem, which connects to the main vine.

The appearance of poison ivy can vary greatly between environments, and even within a single
area. Identification by experienced people is often made difficult by leaf damage, the plant’s leaf-
less condition during winter, and unusual growth forms due to environmental or genetic factors.

Various mnemonic rhymes describe the characteristic appearance of poison ivy:

1. “Leaflets three; let it be” is the best known and most useful cautionary rhyme. It applies
to poison oak, as well as to poison ivy, but other, non-harmful plants have similar leaves.

2. “Hairy vine, no friend of mine.”

3. “Berries white, run in fright” and “Berries white, danger in sight.”

Effects on the Body

A video describing the effects of poison ivy on the body

Blisters from contact with poison ivy

Urushiol-induced contact dermatitis is the allergic reaction caused by poison ivy. In extreme cases,
a reaction can progress to anaphylaxis. Around 15% to 30% of people have no allergic reaction to
urushiol, but most people will have a greater reaction with repeated or more concentrated exposure.

Over 350,000 people are affected by poison ivy annually in the United States.

The pentadecylcatechols of the oleoresin within the sap of poison ivy and related plants causes the
allergic reaction; the plants produce a mixture of pentadecylcatechols, which collectively is called
urushiol. After injury, the sap leaks to the surface of the plant where the urushiol becomes a black-
ish lacquer after contact with oxygen.

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164 Fundamentals of Weed Science

Urushiol binds to the skin on contact, where it causes severe itching that develops into reddish
inflammation or non-coloured bumps, and then blistering. These lesions may be treated with Cal-
amine lotion, Burow’s solution compresses, dedicated commercial poison ivy itch creams, or baths
to relieve discomfort, though recent studies have shown some traditional medicines to be ineffec-
tive. Over-the-counter products to ease itching—or simply oatmeal baths and baking soda—are
now recommended by dermatologists for the treatment of poison ivy. A plant-based remedy cited
to counter urushiol-induced contact dermatitis is jewelweed, and a Jewelweed mash made from
the living plant was effective in reducing poison ivy dermatitis, supporting ethnobotanical use,
while jewelweed extracts had no positive effect in clinical studies. Others argue that prevention of
lesions is easy if one practices effective washing, using plain soap, scrubbing with a washcloth, and
rinsing three times within two to eight hours of exposure.

The oozing fluids released by scratching blisters do not spread the poison. The fluid in the blisters
is produced by the body and it is not urushiol itself. The appearance of a spreading rash indicates
that some areas received more of the poison and reacted sooner than other areas or that contam-
ination is still occurring from contact with objects to which the original poison was spread. Those
affected can unknowingly spread the urushiol inside the house, on phones, door knobs, couches,
counters, desks, and so on, thus in fact repeatedly coming into contact with poison ivy and extend-
ing the length of time of the rash. If this has happened, wipe down the surfaces with bleach or a
commercial urushiol removal agent. The blisters and oozing result from blood vessels that develop
gaps and leak fluid through the skin; if the skin is cooled, the vessels constrict and leak less. If poi-
son ivy is burned and the smoke then inhaled, this rash will appear on the lining of the lungs, caus-
ing extreme pain and possibly fatal respiratory difficulty. If poison ivy is eaten, the mucus lining of
the mouth and digestive tract can be damaged. A poison ivy rash usually develops within a week of
exposure and can last anywhere from one to four weeks, depending on severity and treatment. In
rare cases, poison ivy reactions may require hospitalization.

Urushiol oil can remain active for several years, so handling dead leaves or vines can cause a reac-
tion. In addition, oil transferred from the plant to other objects (such as pet fur) can cause the rash
if it comes into contact with the skin. Clothing, tools, and other objects that have been exposed to
the oil should be washed to prevent further transmission.

People who are sensitive to poison ivy can also experience a similar rash from mangoes. Mangoes
are in the same family (Anacardiaceae) as poison ivy; the sap of the mango tree and skin of man-
goes has a chemical compound similar to urushiol. A related allergenic compound is present in
the raw shells of cashews. Similar reactions have been reported occasionally from contact with the
related Fragrant Sumac (Rhus aromatica) and Japanese lacquer tree. These other plants are also
in the Anacardiaceae family.

Treatment of Poison ivy Rash


Immediate washing with soap and cold water or rubbing alcohol may help prevent a reaction. Hot
water should not be used, as it causes one’s pores to open up and admit the oils from the plant.
During a reaction, calamine lotion or diphenhydramine may help mitigate symptoms. Cortico-
steroids, either applied to the skin or taken by mouth, may be appropriate in extreme cases. An
astringent containing aluminum acetate (such as Burow’s solution) may also provide relief and
soothe the uncomfortable symptoms of the rash.

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Similar-looking Plants
• Virgin’s bower (Clematis virginiana) (also known as Devil’s Darning Needles, Devil’s
Hair, Love Vine, Traveller’s Joy, Virginia Virgin’s Bower, Wild Hops, and Woodbine; syn.
Clematis virginiana L. var. missouriensis (Rydb.) Palmer & Steyermark ) is a vine of the
Ranunculaceae family native to the United States. This plant is a vine that can climb up
to 10–20 ft tall. It grows on the edges of the woods, moist slopes, and fence rows and in
thickets and streambanks. It produces white, fragrant flowers about an inch in diameter
between July and September.

• Box-elder (Acer negundo) saplings have leaves that can look very similar to those of poi-
son ivy, although the symmetry of the plant itself is very different. While box-elders often
have five or seven leaflets, three leaflets are also common, especially on smaller saplings.
The two can be differentiated by observing the placement of the leaves where the leaf stalk
meets the main branch (where the three leaflets are attached). Poison ivy has alternate
leaves, which means the three-leaflet leaves alternate along the main branch. The maple
(which the box-elder is a species of) has opposite leaves; another leaf stalk directly on the
opposite side is characteristic of box-elder.

• Virginia creeper (Parthenocissus quinquefolia) vines can look like poison ivy. The younger
leaves can consist of three leaflets but have a few more serrations along the leaf edge, and
the leaf surface is somewhat wrinkled. However, most Virginia creeper leaves have five
leaflets. Virginia creeper and poison ivy very often grow together, even on the same tree.
Even those who do not get an allergic reaction to poison ivy may be allergic to the oxalate
crystals in Virginia creeper sap.

• Western poison oak (Toxicodendron diversilobum) leaflets also come in threes on the end
of a stem, but each leaflet is shaped somewhat like an oak leaf. Western poison oak grows
only in the western United States and Canada, although many people will refer to poison
ivy as poison oak. This is because poison ivy will grow in either the ivy-like form or the
brushy oak-like form depending on the moisture and brightness of its environment. The
ivy form likes shady areas with only a little sun, tends to climb the trunks of trees, and can
spread rapidly along the ground.

• Poison sumac (Toxicodendron vernix) has compound leaves with 7–15 leaflets. Poison su-
mac never has only three leaflets.

• Kudzu (Pueraria lobata) is a non-toxic edible vine that scrambles extensively over lower
vegetation or grows high into trees. Kudzu is an invasive species in the southern United
States. Like poison ivy, it has three leaflets, but the leaflets are bigger than those of poison
ivy and are pubescent underneath with hairy margins.

• Blackberries and raspberries (Rubus spp.) can resemble poison ivy, with which they may
share territory; however, blackberries and raspberries almost always have thorns on their
stems, whereas poison ivy stems are smooth. Also, the three-leaflet pattern of some black-
berry and raspberry leaves changes as the plant grows: Leaves produced later in the sea-
son have five leaflets rather than three. Blackberries and raspberries have many fine teeth
along the leaf edge, the top surface of their leaves is very wrinkled where the veins are, and

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166 Fundamentals of Weed Science

the bottom of the leaves is light minty-greenish white. Poison ivy is all green. The stem of
poison ivy is brown and cylindrical, while blackberry and raspberry stems can be green, can
be squared in cross-section, and can have prickles. Raspberries and blackberries are never
truly vines; that is, they do not attach to trees to support their stems.

• The thick vines of riverbank grape (Vitis riparia), with no rootlets visible, differ from the
vines of poison ivy, which have so many rootlets that the stem going up a tree looks furry.
Riverbank grape vines are purplish in colour, tend to hang away from their support trees,
and have shreddy bark; poison ivy vines are brown, attached to their support trees, and do
not have shreddy bark.

• Fragrant sumac (Rhus aromatica) has a very similar appearance to poison ivy. While both
species have three leaflets, the center leaflet of poison ivy is on a long stalk, while the center
leaflet of fragrant sumac does not have an obvious stalk. When crushed, fragrant sumac
leaves have a fragrance similar to citrus while poison ivy has little or no distinct fragrance.
Fragrant sumac produces flowers before the leaves in the spring, while poison ivy produces
flowers after the leaves emerge. Flowers and fruits of fragrant sumac are at the end of the
stem, but occur along the middle of the stem of poison ivy. Fragrant sumac fruit ripens to a
deep reddish color and is covered with tiny hairs while poison ivy fruit is smooth and ripens
to a whitish color.

• Hoptree (Ptelea trifoliata) has leaves that are remarkably similar. It is, however, a much
larger plant so confusion is unlikely for any but the smallest specimens. The flowers and
seeds are also easily distinguished from those of poison ivy.

Similar Allergenic Plants


• Toxicodendron rydbergii (Western poison ivy)

• Smodingium argutum (African poison ivy)

• Toxicodendron pubescens (Poison oak – Eastern)

• Toxicodendron diversilobum (Poison oak – Western)

• Toxicodendron vernix (Poison sumac)

• Gluta spp (Rengas tree)

• Toxicodendron vernicifluum (Japanese lacquer tree)

Taraxacum
Taraxacum is a large genus of flowering plants in the family Asteraceae which consists of species
commonly known as dandelion. They are native to Eurasia and North America, but the two com-
monplace species worldwide, T. officinale and T. erythrospermum, were imports from Europe
that now propagate as wildflowers. Both species are edible in their entirety. The common name

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Plants Considered to be Weed 167

dandelion is given to members of the genus. Like other members of the Asteraceae family, they
have very small flowers collected together into a composite flower head. Each single flower in
a head is called a floret. Many Taraxacum species produce seeds asexually by apomixis, where
the seeds are produced without pollination, resulting in offspring that are genetically identical
to the parent plant.

Description
The species of Taraxacum are tap-rooted, perennial, herbaceous plants, native to temperate areas
of the Northern Hemisphere. The genus contains many species which usually (or in the case of
triploids, obligately) reproduce by apomixis, resulting in many local populations and endemism.
In the British Isles alone, 234 microspecies are recognised in 9 loosely defined sections, of which
40 are “probably endemic”.

In general, the leaves are 5–25 cm long or longer, simple, lobed, and form a basal rosette above
the central taproot. The flower heads are yellow to orange coloured, and are open in the daytime,
but closed at night. The heads are borne singly on a hollow stem (scape) that is usually leafless and
rises 1–10 cm or more above the leaves. Stems and leaves exude a white, milky latex when broken.
A rosette may produce several flowering stems at a time. The flower heads are 2–5 cm in diameter
and consist entirely of ray florets. The flower heads mature into spherical seed heads called blow-
balls or clocks (in both British and American English) containing many single-seeded fruits called
achenes. Each achene is attached to a pappus of fine hairs, which enable wind-aided dispersal over
long distances.

The flower head is surrounded by bracts (sometimes mistakenly called sepals) in two series. The
inner bracts are erect until the seeds mature, then flex downward to allow the seeds to disperse.
The outer bracts are often reflexed downward, but remain appressed in plants of the sections Pa-
lustria and Spectabilia. Some species drop the parachute from the achenes; the hair-like para-
chutes are called pappus, and they are modified sepals. Between the pappus and the achene is a
stalk called a beak, which elongates as the fruit matures. The beak breaks off from the achene quite
easily, separating the seed from the parachute.

Seed Dispersal

Segment of pappus fiber showing barbs

A number of species of Taraxacum are seed-dispersed ruderals that rapidly colonize disturbed
soil, especially the common dandelion (T. officinale), which has been introduced over much of the
temperate world. After flowering is finished, the dandelion flower head dries out for a day or two.
The dried petals and stamens drop off, the bracts reflex (curve backwards), and the parachute ball
opens into a full sphere.

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168 Fundamentals of Weed Science

False Dandelions

Hawksbeard flower heads and ripe seeds are sometimes confused with dandelions.

Many similar plants in the Asteraceae family with yellow flowers are sometimes known as false
dandelions. Dandelions are very similar to catsears (Hypochaeris). Both plants carry similar
flowers, which form into windborne seeds. However, dandelion flowers are borne singly on un-
branched, hairless and leafless, hollow stems, while catsear flowering stems are branched, solid,
and carry bracts. Both plants have a basal rosette of leaves and a central taproot. However, the
leaves of dandelions are smooth or glabrous, whereas those of catsears are coarsely hairy.

Early-flowering dandelions may be distinguished from coltsfoot (Tussilago farfara) by their basal
rosette of leaves, their lack of disc florets, and the absence of scales on the flowering stem.

Other plants with superficially similar flowers include hawkweeds (Hieracium) and hawksbeards
(Crepis). These are readily distinguished by branched flowering stems, which are usually hairy and
bear leaves.

Classification
The genus is taxonomically complex, with some botanists dividing the group into about 34 mac-
rospecies, and about 2000 microspecies; about 235 apomictic and polyploid microspecies have
been recorded in Great Britain and Ireland. Some botanists take a much narrower view and only
accept a total of about 60 species.

Selected Species
• Taraxacum albidum, a white-flowering Japanese dandelion
• Taraxacum aphrogenes, Paphos dandelion
• Taraxacum brevicorniculatum, frequently misidentified as Taraxacum kok-saghyz, and
a poor rubber producer
• Taraxacum californicum, the endangered California dandelion
• Taraxacum centrasiaticum, the Xinjiang dandelion
• Taraxacum ceratophorum, northern dandelion

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Plants Considered to be Weed 169

• Taraxacum erythrospermum, often considered a variety of T. laevigatum


• Taraxacum farinosum, Turkish dandelion
• Taraxacum holmboei, Troödos dandelion
• Taraxacum japonicum, Japanese dandelion, no ring of smallish, downward-turned leaves
under the flowerhead
• Taraxacum kok-saghyz, Russian dandelion, which produces rubber
• Taraxacum laevigatum, red-seeded dandelion, achenes reddish brown and leaves deeply
cut throughout length, inner bracts’ tips are hooded
• Taraxacum mirabile
• Taraxacum officinale (syn. T.  officinale subsp. vulgare), common dandelion. Found in
many forms.
• Taraxacum pankhurstianum
• Taraxacum platycarpum, the Korean dandelion

File:T japonicum04.
jpg T. japonicum
File:T albidum01.jpg T. File:Taraxacumcaliforni-
albidum cum.jpg T. californicum

File:Taraxacum laevigatum File:DandelionFlower.jpg


Closeup DehesaBoyaldePuer- T. officinale
tollano.jpg T. laevigatum

Cultivars
• ‘Amélioré à Coeur Plein’ yields an abundant crop without taking up much ground, and
tends to blanch itself naturally, due to its clumping growth habit.
• ‘Broad-leaved’ - The leaves are thick and tender and easily blanched. In rich soils, they can
be up to 60 cm wide. Plants do not go to seed as quickly as French types.
• ‘Vert de Montmagny’ is a large-leaved, vigorous grower, which matures early.

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170 Fundamentals of Weed Science

History
Dandelions are thought to have evolved about 30 million years ago in Eurasia. They have been
used by humans for food and as an herb for much of recorded history.

Names

Leaf resemblance to lion tooth

The Latin name Taraxacum originates in medieval Persian writings on pharmacy. The Persian
scientist Al-Razi around 900 AD wrote “the tarashaquq is like chicory”. The Persian scientist and
philosopher Ibn Sīnā around 1000 AD wrote a book chapter onTaraxacum.

The English name, dandelion, is a corruption of the French dent de lion meaning “lion’s tooth”, re-
ferring to the coarsely toothed leaves. The plant is also known as blowball, cankerwort, doon-head-
clock, witch’s gowan, milk witch, lion’s-tooth, yellow-gowan, Irish daisy, monks-head, priest’s-
crown, and puff-ball; other common names include faceclock, pee-a-bed, wet-a-bed, swine’s
snout, white endive, and wild endive.

The English folk name “piss-a-bed” (and indeed the equivalent contemporary French pissenlit)
refers to the strong diuretic effect of the plant’s roots. In various northeastern Italian dialects, the
plant is known as pisacan (“dog pisses”), because they are found at the side of pavements.

In Swedish, it is called maskros (worm rose) after the small insects (thrips) usually present in the
flowers. In Finnish and Estonian, the names (voikukka, võilill) translate as butter flower, due to
the color of the flower. In Lithuanian, it is known as “Pienė”, meaning “milky”, because of the white
liquid that is produced when the stems are cut. The Welsh (dant-y-llew), German (Löwenzahn),
Norwegian (løvetann) and Spanish (diente de león) names mean the same as the French and the
English names.

Properties
Edibility
Dandelions are found on all continents and have been gathered for food since prehistory, but the
varieties cultivated for consumption are mainly native to Eurasia. A perennial plant, its leaves
will grow back if the taproot is left intact. To make leaves more palatable, they are often blanched
to remove bitterness, or sauteed in the same way as spinach. Dandelion leaves and buds have

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been a part of traditional Kashmiri, Slovenian, Sephardic, Chinese, and Korean cuisines. In
Crete, the leaves of a variety called ‘Mari’, ‘Mariaki’, or ‘Koproradiko’ are eaten by locals, either
raw or boiled, in salads. T. megalorhizon, a species endemic to Crete, is eaten in the same way;
it is found only at high altitudes (1000 to 1600 m) and in fallow sites, and is called pentaramia
or agrioradiko.

The flower petals, along with other ingredients, usually including citrus, are used to make dandeli-
on wine. The ground, roasted roots can be used as a caffeine-free dandelion coffee. Dandelion was
also traditionally used to make the traditional British soft drink dandelion and burdock, and is one
of the ingredients of root beer. Also, dandelions were once delicacies eaten by the Victorian gentry,
mostly in salads and sandwiches.

Dandelion leaves contain abundant vitamins and minerals, especially vitamins A, C, and K, and
are good sources of calcium, potassium, iron, and manganese.

Medicinal Uses
Historically, dandelion was prized for a variety of medicinal properties, and it contains a number
of pharmacologically active compounds. Dandelion is used as a herbal remedy in Europe, North
America, and China. It has been used in herbal medicine to treat infections, bile and liver prob-
lems, and as a diuretic.

Food for Wildlife


Taraxacum seeds are an important food source for certain birds.

Dandelions are also important plants for Northern Hemisphere bees, providing an important source
of nectar and pollen early in the season. Dandelions are used as food plants by the larvae of some spe-
cies of Lepidoptera (butterflies and moths). They are also used as a source of nectar by the pearl-bor-
dered fritillary (Boloria euphrosyne), one of the earliest emerging butterflies in the spring.

Benefits to Gardeners
The dandelion plant is a beneficial weed, with a wide range of uses, and is even a good companion
plant for gardening. Its taproot will bring up nutrients for shallower-rooting plants, and add min-
erals and nitrogen to soil. It is also known to attract pollinating insects and release ethylene gas
which helps fruit to ripen.

Cultural Importance
Four dandelion flowers are the emblem of White Sulphur Springs, West Virginia. The citizens cel-
ebrate spring with an annual Dandelion Festival.

The dandelion is the official flower of the University of Rochester and “Dandelion Yellow” is one
of the school’s official colors. “The Dandelion Yellow” is an official University of Rochester song.

Dangers
Dandelion pollen may cause allergic reactions when eaten, or adverse skin reactions in sensitive

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172 Fundamentals of Weed Science

individuals. Contact dermatitis after handling has also been reported, probably from the latex in
the stems and leaves. Due to its high potassium level, dandelion can also increase the risk of hy-
perkalemia when taken with potassium-sparing diuretics.

As a Noxious Weed
The species T. officinale is listed as a noxious weed in some jurisdictions, and is considered to be a
nuisance in residential and recreational lawns in North America. It is also an important weed in agri-
culture and causes significant economic damage because of its infestation in many crops worldwide.

As Source of Natural Rubber


Dandelions secrete latex when the tissues are cut or broken, yet in the wild type, the latex con-
tent is low and varies greatly. Using modern cultivation methods and optimization techniques,
scientists in the Fraunhofer Institute for Molecular Biology and Applied Ecology (IME) in Ger-
many developed a cultivar that is suitable for commercial production of natural rubber. The la-
tex produced exhibits the same quality as the natural rubber from rubber trees. In collaboration
with Continental Tires, IME is building a pilot facility. As of May 2014, the first prototype test
tires made with blends from dandelion-rubber are scheduled to be tested on public roads over
the next few years.

Plantago Major
Plantago major (broadleaf plantain, white man’s foot, or greater plantain) is a species of flowering
plant in the plantain family Plantaginaceae. The plant is native to most of Europe and northern
and central Asia, but has widely naturalised elsewhere in the world.

Plantago major is one of the most abundant and widely distributed medicinal crops in the world.
A poultice of the leaves can be applied to wounds, stings, and sores in order to facilitate healing
and prevent infection. The active chemical constituents are aucubin (an anti-microbial agent), al-
lantoin (which stimulates cellular growth and tissue regeneration), and mucilage (which reduces
pain and discomfort). Plantain has astringent properties, and a tea made from the leaves can be
ingested to treat diarrhea and soothe raw internal membranes.

Broadleaf plantain is also a highly nutritious wild edible, that is high in calcium and vitamins A, C,
and K. The young, tender leaves can be eaten raw, and the older, stringier leaves can be boiled in
stews and eaten.

Description
Plantago major is an herbaceous perennial plant with a rosette of leaves 15–30 cm in diameter.

Each leaf is oval-shaped, 5–20  cm long and 4–9  cm broad, rarely up to 30  cm long and 17  cm
broad, with an acute apex and a smooth margin; there are five to nine conspicuous veins. The flow-
ers are small, greenish-brown with purple stamens, produced in a dense spike 5–15 cm long on top
of a stem 13–15 cm tall (rarely to 70 cm tall).

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Plantago major is notable for its ability to colonize compacted and disturbed soils,
and to survive repeated trampling.

Plantain is wind-pollinated, and propagates primarily by seeds, which are held on the long, narrow
spikes which rise well above the foliage. Each plant can produce up to 20,000 seeds, which are very
small and oval-shaped, with a bitter taste.

There are three subspecies:


• Plantago major subsp. major.
• Plantago major subsp. intermedia (DC.) Arcang.
• Plantago major subsp. winteri (Wirtg.) W.Ludw.

Ecology

Developing fruits of Plantago major

Plantago major grows in lawns and fields, along roadsides, and in other areas that have been
disturbed by humans. It does particularly well in compacted or disturbed soils. It is believed to be
one of the first plants to reach North America after European colonisation. Reportedly brought to
the Americas by Puritan colonizers, plantain was known among some Native American peoples by
the common name “white man’s footprint”, because it thrived in the disturbed and damaged eco-
systems surrounding European settlements. The ability of plantain to survive frequent trampling
and colonize compacted soils makes it important for soil rehabilitation. Its roots break up hardpan
surfaces, while simultaneously holding together the soil to prevent erosion.

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174 Fundamentals of Weed Science

The seeds of plantain are a common contaminant in cereal grain and other crop seeds. As a result,
it now has a worldwide distribution as a naturalised species and often a weed.

Edibility
The leaves are edible as a salad green when young and tender, but they quickly become tough and
fibrous as they get older. The older leaves can be cooked in stews. The leaves contain calcium and
other minerals, with 100 grams of plantain containing approximately the same amount of vitamin
A as a large carrot. The seeds are so small that they are tedious to gather, but they can be ground
into a flour substitute or extender.

Medicinal Use
Plantain is found all over the world, and is one of the most abundant and accessible medicinal
herbs. It contains many bioactive compounds, including allantoin, aucubin, ursolic acid, flavo-
noids, and asperuloside. Scientific studies have shown that plantain extract has a wide range of
biological effects, including “wound healing activity, anti-inflammatory, analgesic, antioxidant,
weak antibiotic, immuno modulating and antiulcerogenic activity”.

For millennia, poultices of plantain leaves have been applied to wounds, sores, and stings to pro-
mote healing. (The clown Costard cries out for a plantain after cutting his shin in Shakespeare’s
Love’s Labour’s Lost.) The active constituents are the anti-microbial compound aucubin, the cell-
growth promoter allantoin, a large amount of soothing mucilage, flavonoids, caffeic acid deriva-
tives, and alcohols in the wax on the leaf surface. The root of plantain was also traditionally used
to treat wounds, as well as to treat fever and respiratory infections.

Due to its astringent properties, a tea of plantain leaves can be ingested to treat diarrhea or dys-
entery. Due to the high vitamin and mineral content, plantain tea simultaneously replenishes the
nutrients lost as a result of diarrhea. Adding fresh plantain seeds or flower heads to a tea will act
as an effective lubricating and bulking laxative and soothe raw, sore throats.

When ingested, the aucubin in plantain leaves leads to increased uric acid excretion from the kid-
neys, and may be useful in treating gout.

Other Uses

Cultivar ‘Rubrifolia’ of Plantago major

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Plants Considered to be Weed 175

The sinews from the mature plant are very pliable and tough, and can be used in survival situations
to make small cords, fishing line, sutures, or braiding.

Some cultivars are planted as ornamentals in gardens, including ‘Rubrifolia’ with purple leaves,
and ‘Variegata’ with variegated leaves.

Silybum Marianum

Illustration

Leaves

Cotyledons and first leaves

Silybum marianum has other common names include cardus marianus, milk thistle, blessed milk-
thistle, Marian thistle, Mary thistle, Saint Mary’s thistle, Mediterranean milk thistle, variegated
thistle and Scotch thistle. This species is an annual or biennial plant of the Asteraceae family. This
fairly typical thistle has red to purple flowers and shiny pale green leaves with white veins. Origi-
nally a native of Southern Europe through to Asia, it is now found throughout the world.

Description
Milk thistles can grow to be 30 to 200 cm (12 to 79 in) tall, and have an overall conical shape. The

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176 Fundamentals of Weed Science

approximate maximum base diameter is 160 cm (63 in). The stem is grooved and more or less
cottony. The largest specimens have hollow stems.

The leaves are oblong to lanceolate. They are either lobate or pinnate, with spiny edges. They are
hairless, shiny green, with milk-white veins.

The flower heads are 4 to 12 cm long and wide, of red-purple colour. They flower from June to August
in the North or December to February in the Southern Hemisphere (summer through autumn).

The bracts are hairless, with triangular, spine-edged appendages, tipped with a stout yellow spine.

The achenes are black, with a simple long white pappus, surrounded by a yellow basal ring.

Distribution and Habitat


Possibly native near the coast of southeast England, it has been widely introduced outside its nat-
ural range, for example into North America, Iran, Australia and New Zealand where it is consid-
ered an invasive weed. Cultivated fields for the production of raw material for the pharmaceutical
industry exist on a larger scale in Austria (Waldviertel region), Germany, Hungary, Poland, China
and Argentina. In Europe it is sown yearly in March–April. The harvest in two steps (cutting and
threshing) takes place in August, about 2–3 weeks after the flowering.

Exkursionsflora Fuer Kreta by Jahn & Schoenfelder (1995, page 326) states that the distribution is
mediterranean-near east. They quote it as a native plant of Crete, Greece.

Milk thistle (Silybum marianum) is a thorny plant presenting decorative leaves with a white pat-
tern of veins and purple flower heads. The plant originates from mountains of the Mediterranean
region, where it forms scrub on a rocky base.

The plant is sometimes also used as a decorative element in gardens, and its dried flower heads
may be used for the decoration of dry bouquets.

The “giant thistle of the Pampas” reported by Darwin in the Voyage of the Beagle is thought by
some to be Silybum marianum.

FMIB 47258 Giant Thistle of Pampas

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Plants Considered to be Weed 177

Chemistry
Traditional milk thistle extract is made from the seeds, which contain approximately 4–6% sily-
marin. The extract consists of about 65–80% silymarin (a flavonolignan complex) and 20–35%
fatty acids, including linoleic acid. Silymarin is a complex mixture of polyphenolic molecules, in-
cluding seven closely related flavonolignans (silybin A, silybin B, isosilybin A, isosilybin B, sily-
christin, isosilychristin, silydianin) and one flavonoid (taxifolin). Silibinin, a semipurified fraction
of silymarin, is primarily a mixture of 2 diastereoisomers, silybin A and silybin B, in a roughly 1:1
ratio.

Medicinal Use
Clinical Trials
Milk thistle has been researched for a number of purposes including treatment of liver disease, and
cancer; however, clinical studies are largely heterogeneous and contradictory.

In trials, silymarin has typically been administered in amounts ranging from 420–480 mg per day
in two to three divided doses. However, higher doses have been studied, such as 600 mg daily in
the treatment of type II diabetes (with significant results), and 600 or 1200 mg daily in patients
chronically infected with hepatitis C virus (without significant results). An optimal dosage for milk
thistle preparations has not been established.

Herbal Medicinal Research


Silybum marianum is used in traditional Chinese medicine to clear heat and relieve toxic ma-
terial, to soothe the liver and to promote bile flow. Though its efficacy in treating diseases is still
unknown, Silybum marianum is sometimes prescribed by herbalists to help treat liver diseases
(cirrhosis, jaundice and hepatitis). Both in vitro and animal research suggest that Silibinin (syn.
silybin, sylimarin I) may have hepatoprotective (antihepatotoxic) properties that protect liver cells
against toxins.

A 2000 study of such claims by the AHRQ concluded that “clinical efficacy of milk thistle is not
clearly established”. A 2005 Cochrane Review considered thirteen randomized clinical trials which
assessed milk thistle in 915 patients with alcoholic and/or hepatitis B or C virus liver diseases.
They question the beneficial effects of milk thistle for patients with alcoholic and/or hepatitis B or
C virus liver diseases and highlight the lack of high-quality evidence to support this intervention.
Cochrane concluded that more good-quality randomized clinical trials on milk thistle versus pla-
cebo are needed.

Cancer Research UK say that milk thistle is promoted on the internet for its claimed ability to slow
certain kinds of cancer, but that there is no good evidence in support of these claims.

Safety
Milk thistle extracts are known to be safe and well-tolerated. Milk thistle supplements, however,
were measured to have the highest mycotoxin concentrations of up to 37 mg/kg when compared
amongst various plant-based dietary supplements.

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178 Fundamentals of Weed Science

Use as Food
Milk thistle has also been known to be used as food. The roots can be eaten raw or boiled and but-
tered or par-boiled and roasted. The young shoots in spring can be cut down to the root and boiled
and buttered. The spiny bracts on the flower head were eaten in the past like globe artichoke, and
the stems (after peeling) can be soaked overnight to remove bitterness and then stewed. The leaves
can be trimmed of prickles and boiled as a spinach substitute or they can also be added raw to
salads.

Animal Toxicity
Because of potassium nitrate content, the plant has been found to be toxic to cattle and sheep.
When potassium nitrate is eaten by ruminants, the bacteria in the animal’s stomach breaks the
chemical down, producing nitrite ions. Nitrite ions then combine with hemoglobin to produce
methaemoglobin, blocking the transport of oxygen. The result is a form of oxygen deprivation.

Trifolium Repens

Trifolium repens, the white clover (also known as Dutch clover, Ladino clover, or Ladino), is a
herbaceous perennial plant in the bean family Fabaceae. It is native to Europe and central Asia.
One of the most widely cultivated types of clover, it has been widely introduced worldwide as a
forage crop, and is now also common in most grassy areas (lawns and gardens) of North America
and New Zealand. The species includes varieties often classed as small, intermediate and large,
according to height, which reflects petiole length. The term “white clover” is applied to the species
in general, “Dutch clover” is often applied to intermediate varieties (but sometimes to smaller va-
rieties), and “ladino clover” is applied to large varieties.

Name
The genus name, Trifolium, derives from the Latin tres, “three”, and folium, “leaf”, so called from
the characteristic form of the leaf, which usually but not always has three leaflets (trifoliate); hence
the popular name “trefoil”. The species name, repens, is Latin for “creeping”.

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Plants Considered to be Weed 179

Description
It is a herbaceous, perennial plant. It is low growing, with heads of whitish flowers, often with
a tinge of pink or cream that may come on with the aging of the plant. The heads are generally
1.5–2 centimetres (0.6–0.8  in) wide, and are at the end of 7-cm (2.8-in) peduncles or flower
stalks. The flowers are mostly visited by bumblebees and often by honey bees. The leaves, are tri-
foliolate, smooth, elliptic to egg-shaped and long-petioled and usually with light or dark mark-
ings. The stems function as stolons, so white clover often forms mats, with the stems creeping
as much as 18 cm (7.1 in) a year, and rooting at the nodes. The leaves form the symbol known as
shamrock.

Cultivation and Uses


Forage
White clover has been described as the most important forage legume of the temperate zones.
Symbiotic nitrogen fixation (up to 545 kg N per hectare per year, although usually much less, e.g.
about 110 to 170 kg N per hectare per year) in root nodules of white clover obviates synthetic nitro-
gen fertilizer use for maintaining productivity on much temperate zone pasture land. White clover
is commonly grown in mixtures with forage grasses, e.g. perennial ryegrass (Lolium perenne),
Such mixtures can not only optimize livestock production, but can also reduce the bloat risk to
livestock that can be associated with excessive white clover in pastures. Such species mixtures
also tend to avoid issues that could otherwise be associated with cyanogenic glycoside (linamarin
and lotaustralin) intake on pure or nearly pure stands of some white clover varieties. Howev-
er, problems do not inevitably arise with grazing on monocultures of white clover, and superior
ruminant production is sometimes achieved on white clover monocultures managed to optimize
sward height. Formononetin and biochanin A play a role in arbuscular mycorrhiza formation on
white clover roots, and foliar disease can stimulate production of estrogenic coumestans in white
clover. However while there have been a few reports of phytoestrogenic effects of white clover on
grazing ruminants, these have been far less common than such reports regarding some varieties of
subterranean and red clover. Among forage plants, some white clover varieties tend to be favored
by rather close grazing, because of their stoloniferous habit, which can contribute to competitive
advantage. T. repens is also considered as a folk medicine in India against intesinal helminthic
worms and an experimental in vivo study validated that the aerial shoots of T. repens bear signif-
icant anticestodal properties.

Companion Planting, Green Manure, and Cover Crops


White clover grows well as a companion plant among turf grasses, grain crops, pasture grasses,
and vegetable rows. White clover can tolerate close mowing and grazing, and it can grow on many
different types and pHs of soil (although it prefers clay soils). As a leguminous and hardy plant,
it is considered to be a beneficial component of natural or organic pasture management and lawn
care due to its ability to fix nitrogen and out-compete weeds. Natural nitrogen fixing reduces leach-
ing from the soil and by maintaining soil health can reduce the incidence of some lawn diseases
that are enhanced by the availability of synthetic fertilizer. For these reasons, it is often used as a
green manure and cover crop.

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180 Fundamentals of Weed Science

Culinary Uses
Besides making an excellent forage crop for livestock, clovers are a valuable survival food: they
are high in proteins, widespread, and abundant. The fresh plants have been used for centuries as
additives to salads and other meals consisting of leafy vegetables. They are not easy for humans to
digest raw, however, but this is easily fixed by boiling the harvested plants for 5–10 minutes.

White clover with four leaves.

Four leaf Trifolium repens, in its natural setting. Three-leaf shamrocks can be seen

As an Invasive Weed
Before the introduction of broad-leaf herbicides, white clover was more often added to lawn seed
mixes than it is today, as it is able to grow and provide green cover in poorer soils where turfgrasses
do not perform well. Many people consider clover a weed when growing in lawns, in part because
the flowers are attractive to bees and thus could create a danger for people with bare feet.

White clover is the only known plant on which the caterpillars of the Coleophoridae case-bearer
moth Coleophora mayrella feed.

In Britain, a high abundance of white clover is generally associated with species-poor, agri-
culturally improved grassland habitats, as it out-competes the more rare plants and grasses
especially in fertile soils, and has often been added as part of reseeding. Agri-environment
schemes, such as the Countryside Stewardship Scheme, Environmentally Sensitive Areas
Scheme, and Environmental Stewardship, give funding to species-rich grasslands that are rel-
atively infertile and do not generally have an abundance of white clover. However, white clover
does have value as a pollen and nectar source particularly in intensively farmed areas or amen-
ity grasslands where there are few other flowers, and it can be found naturally at low levels in
species-rich grasslands.

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Plants Considered to be Weed 181

Varieties + Subspecies
• Trifolium repens subsp. macrorrhizum (Boiss.) Ponert
• Trifolium repens var. nevadense (Boiss.) C.Vicioso
• Trifolium repens var. ochranthum K.Maly
• Trifolium repens var. orbelicum (Velen.) Fritsch
• Trifolium repens var. orphanideum (Boiss.) Boiss.
• Trifolium repens subsp. prostratum Nyman

Striga
Striga, commonly known as witchweed or witchers weed, is a genus of parasitic plants that occur
naturally in parts of Africa, Asia, and Australia. It is in the family Orobanchaceae. Some species are
serious pathogens of crop cereals, with the greatest effects being in savanna agriculture in Africa.
It also causes considerable crop losses in other regions, including other tropical and subtropical
crops in its native range and in the Americas.

Description
Witchweeds are characterized by bright-green stems and leaves and small, brightly colored and at-
tractive flowers. They are obligate hemiparasites of roots and require a living host for germination
and initial development, though they can then survive on their own.

The genus is classified in the family Orobanchaceae, although older classifications place it in the
Scrophulariaceae.

The number of species is not certain, but exceeds 40 by some counts.

Hosts and Symptoms


Although most species of Striga are not pathogens that affect human agriculture, some species
have devastating effects upon crops, particularly those planted by subsistence farmers. Crops most
commonly affected are corn, sorghum, rice and sugarcane. Three species cause the most damage:
Striga asiatica, S. gesnerioides, and S. hermonthica.

Witchweed parasitizes maize, millet, sorghum, sugarcane, rice, legumes, and a range of weedy
grasses. It is capable of significantly reducing yields, in some cases wiping out the entire crop.

Host plant symptoms, such as stunting, wilting, and chlorosis, are similar to those seen from se-
vere drought damage, nutrient deficiency, and vascular disease.

Lifecycle
Each plant is capable of producing between 90,000 and 500,000 seeds, which may remain via-

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182 Fundamentals of Weed Science

ble in the soil for over 10 years. Most seeds produced are not viable. An annual plant, witchweed
overwinters in the seed stage. Its seeds germinate in the presence of host root exudate, and devel-
op haustoria which penetrate host root cells. Host root exudate contain strigolactones, signaling
molecules that promote striga seed germination. A bell-like swell forms where the parasitic roots
attach to the roots of the host. The pathogen colonizes underground, where it may spend the next
four to seven weeks before emergence, when it rapidly flowers and produces seeds. Witchweed
seeds spread easily by wind, water, and soil via animal vectors. The chief means of dispersal, how-
ever, is through human interaction, by means of machinery, tools, and clothing.

Photo of plant roots with connected Striga plant

Haustorium Development
Once germination is stimulated, the Striga seed sends out an initial root to probe the soil for the
host root. The initial root secretes an oxidizing enzyme that digests the host root surface, releasing
quinones. If the quinone product is within the appropriate concentrations, a haustorium will develop
from the initial root. The haustorium grows toward the host root until it makes contact with the root
surface, establishing parasitic contact in relatively short order. Within 12 hours of initial haustorium
growth, the haustorium recognizes the host root and begins rapid cell division and elongation. The
haustorium forms a wedge shape and uses mechanical force and chemical digestion to penetrate the
host root, pushing the host cells out of the way. Within 48–72 hours, the haustorium has penetrated
the host root cortex. Finger-like structures on the haustorium, called osculum, penetrate the host xy-
lem through pits in the membrane. The osculum then swell to secure their position within the xylem
membrane. Striga sieve tubes develop along with the osculum. Shortly after the host xylem is pen-
etrated, Striga sieve tubes develop and approach the host phloem within eight cells. This eight cell
layer allows for nonspecific nutrient transport from the host to the Striga seedling. Within 24 hours
after tapping the host xylem and phloem, the Striga cotyledons emerge from the seed.

Environment
Temperatures ranging from 30 to 35 °C (86 to 95 °F) in a moist environment are ideal for germination.
Witchweed will not develop in temperatures below 20 °C (68 °F). Agricultural land with light soil and
low nitrogen levels tend to favor its development. Still, witchweed has demonstrated a wide tolerance

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Plants Considered to be Weed 183

for soil types if soil temperatures are favorably high. Seeds have been shown to survive in frozen soil of
temperatures as low as −15 °C (5 °F), attesting to their aptitude as overwintering structures.

Soil temperature, air temperature, photoperiod, soil type, and soil nutrient and moisture levels do
not greatly deter the development of witchweed. These findings suggest, though it has been limited
to the Carolinas in the United States, that the pathogen would successfully infect the massive corn
crops of the Midwest.

Management
Management of witchweed is difficult because the majority of its life cycle takes place below ground.
If it is not detected before emergence, it is too late to reduce crop loss. To prevent witchweed from
spreading it is necessary to plant uncontaminated seeds and clean soil and plant debris off of
machinery, shoes, clothing, and tools before entering fields. If populations are low, hand weeding
before seeds are produced is an option.

Striga in the United States has been controlled through the use of several management strategies,
including quarantines imposed on affected areas, control of movement of farm equipment be-
tween infected and uninfected areas, herbicide application, and imposed “suicidal germination”.
For the latter, in fields not yet planted in crops, seeds present in the soil are induced to germinate
by injecting ethylene gas, which mimics the natural physiological response tied to host recogni-
tion. Because no host roots are available, the seedlings die. Unfortunately, each Striga plant can
produce tens of thousands of tiny seeds, which can remain dormant in the soil for many years.
Thus, such treatments do not remove all seeds from the soil. Moreover, this method is expensive
and not generally available to many farmers in developing nations of Africa and Asia.

Another method called trap cropping involves planting a species in an infested field that will induce
the Striga seeds to germinate but will not support attachment of the parasite. This method has been
used in sorghum plantations by planting Celosia argentea between the sorghum. Planting silverleaf
(Desmodium uncinatum) inhibits striga seed germination and has worked effectively intercropped
with maize. Cotton, sunflower, linseed, and witchweed-resistant maize are also effective trap crops.

Increasing nitrogen levels in the soil, growing tolerant varieties,and trap-cropping, and planting
susceptible crops harvested before witchweed seed is produced, can also be used. Coating maize
seeds with fungi or a herbicide also appears to be a promising approach.

Several sorghum varieties have high levels of resistance in local conditions, including ‘N-13’, ‘Frami-
da’, and ‘Serena’. ‘Buruma’, ‘Shibe’, ‘Okoa’ and ‘Serere 17’ millet cultivars are considered to be resis-
tant in Tanzania. Some corn varieties show partial resistance to witchweed, including ‘Katumani’ in
Kenya. In a number of rice cultivars, including some cultivars of NERICA (New Rice for Africa), ef-
fective pre- and post- attachment resistance mechanisms have been identified. Some crop cultivars,
especially corn, are resistant to herbicides that kill witchweed when the haustoria attach to the crop.
‘Strigaway’ maize has been shown to reduce the seed bank of striga by 30% in two seasons.

Importance
Maize, sorghum, and sugarcane crops affected by witchweed in the United States have an estimat-

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184 Fundamentals of Weed Science

ed value well over $20 billion. Furthermore, witchweed is capable of wiping out an entire crop. In
fact, it is so prolific that in 1957 Congress allocated money in an attempt to eradicate witchweed.
Thus, the Animal and Plant Health Inspection Service (APHIS) of the U.S. Department of Agri-
culture established a research station and control methods. Through infestation mapping, quar-
antine, and control activities such as contaminated seed destruction, the acreage parasitized by
witchweed has been reduced 99% since its discovery in the United States. APHIS has even offered
cash rewards those who identify and report the weed, and encourages landowners to check their
own acreage.

Parasitizing important economic plants, witchweed is one of the most destructive pathogens in Af-
rica. In fact, witchweed affects 40% of Africa’s arable savanna region, resulting in up to $13 billion
lost every year. Striga affects 40 million hectares (98,842,153 acres) of crops in sub-Saharan Africa
alone. The witchweed infestation is so bad in parts of Africa, some farmers have to relocate every
few years. Furthermore, the majority of crops in Africa are grown by subsistence farmers who can-
not afford expensive witchweed controls, who therefore suffer much as a result of this pathogen.

Common Species

Photo of a striga plant (witchweed)

Striga bilabiata

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Plants Considered to be Weed 185

• Striga asiatica has a very wide geographic distribution, from Africa through southern and
eastern Asia to Australia. Since the 1950s, it is also known from the United States. This
introduction, likely a result of human activity, resulted in an infestation of corn (maize)
across many counties in North and South Carolina. The United States Department of Ag-
riculture and state agencies imposed a quarantine on this area to control its spread - a
process that was apparently successful.

• Striga gesnerioides, cowpea witchweed, as its name implies, is a parasite of cowpea (Vigna
unguiculata), which is not a grass, but a member of the legume family (Fabaceae or Legu-
minosae). This species was also accidentally introduced into Florida in the United States,
where it was found parasitizing Indigofera hirsuta (hairy indigo, another legume).

• Striga hermonthica (purple witchweed) is also a parasite that affects grasses, particularly
sorghum and pearl millet in sub-Saharan Africa (Senegal to Ethiopia, Democratic Republic
of Congo and Tanzania, Angola, Namibia).

• Striga aequinoctialis West Africa

• Striga angolensis Angola

• Striga angustifolia East Africa, Asia, Indonesia

• Striga asiatica (Asiatic witchweed) Africa, Arabian peninsula, India, Burma, China, In-
donesia, the Philippines, Malaysia, New Guinea, Australia (introduced?), USA (intro-
duced)

• Striga aspera Africa

• Striga bilabiata Africa

• Striga brachycalyx Africa.

• Striga chrysantha Central Africa

• Striga dalzielii West Africa

• Striga elegans Angola, Malawi, South Africa, Zimbabwe

• Striga forbesii Africa, Madagascar

• Striga gastonii Chad, Central African Republic

• Striga gesnerioides (cowpea witchweed) Africa, Arabian peninsula, India, USA (intro-
duced)

• Striga gracillima Tanzania

• Striga hallaei Gabon, Democratic Republic of Congo

• Striga hirsuta Madagascar

• Striga indica India

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186 Fundamentals of Weed Science

• Striga junodii South Africa, Mozambique

• Striga klingii West Africa, Nigeria, Ghana, Cameroon, Togo

• Striga latericea East Africa, Ethiopia, Somalia

• Striga lepidagathidis Senegal, Guinea, Guinea Bissau

• Striga lutea Sudan, Ethiopia

• Striga macrantha West Africa, Nigeria, Ivory Coast, Togo

• Striga passargei West and Central Africa, Arabian peninsula

• Striga pinnatifida Ethiopia

• Striga primuloides Ivory Coast, Nigeria

• Striga pubiflora Somalia

• Striga yemenica Ethiopia

References
• National Research Council (2006-10-27). “Amaranth”. Lost Crops of Africa: Volume II: Vegetables. Lost Crops
of Africa. 2. National Academies Press. ISBN 978-0-309-10333-6. OCLC 34344933. Retrieved 2008-07-15.

• Goode, P. M. (1989). Edible plants of Uganda. Food and Agriculture Organization of the United Nations. pp.
25–6. ISBN 9789251027134.

• Donald G. Crosby (2004). The Poisoned Weed: Plants Toxic to Skin. Oxford University Press. pp. 32–. ISBN
978-0-19-515548-8.

• Neil L. Jennings (2010). In Plain Sight: Exploring the Natural Wonders of Southern Alberta. Rocky Mountain
Books Ltd. pp. 105–. ISBN 978-1-897522-78-3.

• Robert L. Rietschel; Joseph F. Fowler; Alexander A. Fisher (2008). Fisher’s contact dermatitis. PMPH-USA.
pp. 408–. ISBN 978-1-55009-378-0. Retrieved 26 July 2010.

• Editors of Prevention (2 March 2010). The Doctors Book of Home Remedies: Quick Fixes, Clever Techniques,
and Uncommon Cures to Get You Feeling Better Fast. Rodale. pp. 488–. ISBN 978-1-60529-866-5.

• Robert Alan Lewis (1998). Lewis’ dictionary of toxicology. CRC Press. pp. 901–. ISBN 978-1-56670-223-2.
Retrieved 18 August 2010.

• Stace, C.A. (2010). New flora of the British isles (Third ed.). Cambridge, U.K.: Cambridge University Press. p.
712. ISBN 9780521707725.

• Blamey, M.; Fitter, R.; Fitter, A (2003). Wild flowers of Britain and Ireland: The Complete Guide to the British
and Irish Flora. London: A & C Black. p. 274. ISBN 978-1408179505.

• Richards, A.J. (1997). Dandelions of Great Britain and Ireland (Handbooks for Field Identification). BSBI Pub-
lications. p. 330. ISBN 978-0-901158-25-3.

• McGee, Harold (2004). “A survey of common vegetables”. On Food and Cooking: the science and lore of the
kitchen. New York: Scribner. p. 320. ISBN 0-684-80001-2.

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5
Weed Control: Methods and Techniques
Controlling or stopping unwanted plants is termed as weed control. It is very important for agri-
culture as weed can compete with domesticated plants. The methods and techniques discussed
in this section are aquatic weed harvester, soil steam sterilization, stale seed bed, drip irrigation,
bush regeneration and herbicide. The aspects elucidated in the chapter are of vital importance,
and provides a better understanding of weed control.

Weed Control
Weed control is the botanical component of pest control, which attempts to stop weeds, especially
noxious or injurious weeds, from competing with domesticated plants and livestock. Weed control
is important in agriculture. Many strategies have been developed in order to contain these plants.
Methods include hand cultivation with hoes, powered cultivation with cultivators, smothering with
mulch, lethal wilting with high heat, burning, and chemical attack with herbicides (weed killers).

A plant is often termed a “weed” when it has one or more of the following characteristics:

• Little or no recognized value (as in medicinal, material, nutritional or energy)

• Rapid growth and/or ease of germination

• Competitive with crops for space, light, water and nutrients

The definition of a weed is completely context-dependent. To one person, one plant may be a weed,
and to another person it may be a desirable plant. In one place, a plant may be viewed as a weed,
whereas in another place, the same plant may be desirable.

Introduction
Weeds compete with productive crops or pasture, ultimately converting productive land into un-
usable scrub. Weeds can be poisonous, distasteful, produce burrs, thorns or otherwise interfere
with the use and management of desirable plants by contaminating harvests or interfering with
livestock.

Weeds compete with crops for space, nutrients, water and light. Smaller, slower growing seedlings
are more susceptible than those that are larger and more vigorous. Onions are one of the most
vulnerable, because they are slow to germinate and produce slender, upright stems. By contrast
broad beans produce large seedlings and suffer far fewer effects other than during periods of water
shortage at the crucial time when the pods are filling out. Transplanted crops raised in sterile soil
or potting compost gain a head start over germinating weeds.

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188 Fundamentals of Weed Science

Weeds also vary in their competitive abilities and according to conditions and season. Tall-grow-
ing vigorous weeds such as fat hen (Chenopodium album) can have the most pronounced effects
on adjacent crops, although seedlings of fat hen that appear in late summer produce only small
plants. Chickweed (Stellaria media), a low growing plant, can happily co-exist with a tall crop
during the summer, but plants that have overwintered will grow rapidly in early spring and may
swamp crops such as onions or spring greens.

The presence of weeds does not necessarily mean that they are damaging a crop, especially during
the early growth stages when both weeds and crops can grow without interference. However, as
growth proceeds they each begin to require greater amounts of water and nutrients. Estimates
suggest that weed and crop can co-exist harmoniously for around three weeks before competition
becomes significant. One study found that after competition had started, the final yield of onion
bulbs was reduced at almost 4% per day.

Perennial weeds with bulbils, such as lesser celandine and oxalis, or with persistent underground
stems such as couch grass (Agropyron repens) or creeping buttercup (Ranunculus repens) store
reserves of food, and are thus able to grow faster and with more vigour than their annual counter-
parts. Some perennials such as couch grass exude allelopathic chemicals that inhibit the growth of
other nearby plants.

Weeds can also host pests and diseases that can spread to cultivated crops. Charlock and Shep-
herd’s purse may carry clubroot, eelworm can be harboured by chickweed, fat hen and shepherd’s
purse, while the cucumber mosaic virus, which can devastate the cucurbit family, is carried by a
range of different weeds including chickweed and groundsel.

Insect pests often do not attack weeds. However pests such as cutworms may first attack weeds
then move on to cultivated crops.

Some plants are considered weeds by some farmers and crops by others. Charlock, a common
weed in the southeastern US, are weeds according to row crop growers, but are valued by bee-
keepers, who seek out places where it blooms all winter, thus providing pollen for honeybees
and other pollinators. Its bloom resists all but a very hard freeze, and recovers once the freeze
ends.

Weed Propagation
Seeds
Annual and biennial weeds such as chickweed, annual meadow grass, shepherd’s purse,
groundsel, fat hen, cleaver, speedwell and hairy bittercress propagate themselves by seeding.
Many produce huge numbers of seed several times a season, some all year round. Groundsel
can produce 1000 seed, and can continue right through a mild winter, whilst Scentless May-
weedproduces over 30,000 seeds per plant. Not all of these will germinate at once, but over
several seasons, lying dormant in the soil sometimes for years until exposed to light. Poppy
seed can survive 80–100 years, dock 50 or more. There can be many thousands of seeds in a
square foot or square metre of ground, thus and soil disturbance will produce a flush of fresh
weed seedlings.

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Weed Control: Methods and Techniques 189

Subsurface/Surface
The most persistent perennials spread by underground creeping rhizomes that can regrow from
a tiny fragment. These include couch grass, bindweed, ground elder, nettles, rosebay willow herb,
Japanese knotweed, horsetail and bracken, as well as creeping thistle, whose tap roots can put out
lateral roots. Other perennials put out runners that spread along the soil surface. As they creep
they set down roots, enabling them to colonise bare ground with great rapidity. These include
creeping buttercup and ground ivy. Yet another group of perennials propagate by stolons- stems
that arch back into the ground to reroot. The most familiar of these is the bramble.

Methods
Weed control plans typically consist of many methods which are divided into biological, chemical,
cultural, and physical/mechanical control.

Pesticide-free thermic weed control with a weed burner on a potato field in Dithmarschen

Physical/Mechanical Methods
Coverings
In domestic gardens, methods of weed control include covering an area of ground with a material
that creates a hostile environment for weed growth, known as a weed mat.

Several layers of wet newspaper prevent light from reaching plants beneath, which kills them. Dai-
ly saturating the newspaper with water plant decomposition. After several weeks, all germinating
weed seeds are dead.

In the case of black plastic, the greenhouse effect kills the plants. Although the black plastic sheet is
effective at preventing weeds that it covers, it is difficult to achieve complete coverage. Eradicating
persistent perennials may require the sheets to be left in place for at least two seasons.

Some plants are said to produce root exudates that suppress herbaceous weeds. Tagetes minuta is
claimed to be effective against couch and ground elder, whilst a border of comfrey is also said to act

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190 Fundamentals of Weed Science

as a barrier against the invasion of some weeds including couch. A 5–10 centimetres (2.0–3.9 in)}
layer of wood chip mulch prevents most weeds from sprouting.

Gravel can serve as an inorganic mulch.

Irrigation is sometimes used as a weed control measure such as in the case of paddy fields to kill
any plant other than the water-tolerant rice crop.

Manual Removal

Weeds are removed manually in large parts of India.

Many gardeners still remove weeds by manually pulling them out of the ground, making sure to
include the roots that would otherwise allow them to resprout.

Hoeing off weed leaves and stems as soon as they appear can eventually weaken and kill perenni-
als, although this will require persistence in the case of plants such as bindweed. Nettle infesta-
tions can be tackled by cutting back at least three times a year, repeated over a three-year period.
Bramble can be dealt with in a similar way.

Tillage
Ploughing includes tilling of soil, intercultural ploughing and summer ploughing. Ploughing up-
roots weeds, causing them to die. In summer ploughing is done during deep summers. Summer
ploughing also helps in killing pests.

Mechanical tilling can remove weeds around crop plants at various points in the growing process.

Thermal
Several thermal methods can control weeds.

Flame weeders use a flame several centimeters away from the weeds to give them a sudden and
severe heating. The goal of flame weeding is not necessarily burning the plant, but rather causing a
lethal wilting by denaturing proteins in the weed. Similarly, hot air weeders can heat up the seeds
to the point of destroying them. Flame weeders can be combined with techniques such as stale
seedbeds (preparing and watering the seedbed early, then killing the nascent crop of weeds that

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Weed Control: Methods and Techniques 191

springs up from it, then sowing the crop seeds) and preemergence flaming (doing a flame pass
against weed seedlings after the sowing of the crop seeds but before those seedlings emerge from
the soil—a span of time that can be days or weeks).

Hot foam (foamstream) causes the cell walls to rupture, killing the plant. Weed burners heat up
soil quickly and destroy superficial parts of the plants. Weed seeds are often heat resistant and
even react with an increase of growth on dry heat.

Since the 19th century soil steam sterilization has been used to clean weeds completely from soil.
Several research results confirm the high effectiveness of humid heat against weeds and its seeds.

Soil solarization in some circumstances is very effective at eliminating weeds while maintaining
grass. Planted grass tends to have a higher heat/humidity tolerance than unwanted weeds.

Seed Targeting
In 1998, the Australian Herbicide Resistance Initiative (AHRI), debuted. gathered fifteen scientists
and technical staff members to conduct field surveys, collect seeds, test for resistance and study the
biochemical and genetic mechanisms of resistance. A collaboration with DuPont led to a mandatory
herbicide labeling program, in which each mode of action is clearly identified by a letter of the alphabet.

The key innovation of the AHRI approach has been to focus on weed seeds. Ryegrass seeds last
only a few years in soil, so if farmers can prevent new seeds from arriving, the number of sprouts
will shrink each year. Until the new approach farmers were unintentionally helping the seeds.
Their combines loosen ryegrass seeds from their stalks and spread them over the fields. In the
mid-1980s, a few farmers hitched covered trailers, called “chaff carts”, behind their combines to
catch the chaff and weed seeds. The collected material is then burned.

An alternative is to concentrate the seeds into a half-meter-wide strip called a windrow and burn
the windrows after the harvest, destroying the seeds. Since 2003, windrow burning has been ad-
opted by about 70% of farmers in Western Australia.

Yet another approach is the Harrington Seed Destructor, which is an adaptation of a coal pulver-
izing cage mill that uses steel bars whirling at up to 1500 rpm. It keeps all the organic material in
the field and does not involve combustion, but kills 95% of seeds.

Cultural Methods
Stale Seed Bed
Another manual technique is the ‘stale seed bed’, which involves cultivating the soil, then leaving
it fallow for a week or so. When the initial weeds sprout, the grower lightly hoes them away before
planting the desired crop. However, even a freshly cleared bed is susceptible to airborne seed from
elsewhere, as well as seed carried by passing animals on their fur, or from imported manure.

Buried Drip Irrigation


Buried drip irrigation involves burying drip tape in the subsurface near the planting bed, thereby

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192 Fundamentals of Weed Science

limiting weeds access to water while also allowing crops to obtain moisture. It is most effective
during dry periods.

Crop Rotation
Rotating crops with ones that kill weeds by choking them out, such as hemp, Mucuna pruriens,
and other crops, can be a very effective method of weed control. It is a way to avoid the use of her-
bicides, and to gain the benefits of crop rotation.

Biological Methods
A biological weed control regiment can consist of biological control agents, bioherbicides, use of
grazing animals, and protection of natural predators.

Animal Grazing
Companies using goats to control and eradicate leafy spurge, knapweed, and other toxic weeds
have sprouted across the American West.

Chemical Methods
“Organic” Approaches

Weed control, circa 1930-40s

A mechanical weed control device: the diagonal weeder

Organic weed control involves anything other than applying manufactured chemicals. Typically a
combination of methods are used to achieve satisfactory control.

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Weed Control: Methods and Techniques 193

Sulfur in some circumstances is accepted within British Soil Association standards.

Herbicides
The above described methods of weed control use no or very limited chemical inputs. They are
preferred by organic gardeners or organic farmers.

However weed control can also be achieved by the use of herbicides. Selective herbicides kill cer-
tain targets while leaving the desired crop relatively unharmed. Some of these act by interfering
with the growth of the weed and are often based on plant hormones. Herbicides are generally
classified as follows:

• Contact herbicides destroy only plant tissue that contacts the herbicide. Generally, these
are the fastest-acting herbicides. They are ineffective on perennial plants that can re-grow
from roots or tubers.

• Systemic herbicides are foliar-applied and move through the plant where they destroy a
greater amount of tissue. Glyphosate is currently the most used systemic herbicide.

• Soil-borne herbicides are applied to the soil and are taken up by the roots of the target
plant.

• Pre-emergent herbicides are applied to the soil and prevent germination or early growth
of weed seeds.

In agriculture large scale and systematic procedures are usually required, often by machines, such
as large liquid herbicide ‘floater’ sprayers, or aerial application.

Bradley Method
Which uses ecological processes to do much of the work. Perennial weeds also propagate by seed-
ing; the airborne seed of the dandelion and the rose-bay willow herb parachute far and wide. Dan-
delion and dock also put down deep tap roots, which, although they do not spread underground,
are able to regrow from any remaining piece left in the ground.

Hybrid
One method of maintaining the effectiveness of individual strategies is to combine them with
others that work in complete different ways. Thus seed targeting has been combined with
herbicides. In Australia seed management has been effectively combined with trifluralin and
clethodim.

Resistance
Resistance occurs when a target adapts to circumvent a particular control strategy. It affects not
only weed control,but antibiotics, insect control and other domains. In agriculture is mostly con-
sidered in reference to pesticides, but can defeat other strategies, e.g., when a target species be-
comes more drought tolerant via selection pressure.

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194 Fundamentals of Weed Science

Farming Practices
Herbicide resistance recently became a critical problem as many Australian sheep farmers switched
to exclusively growing wheat in their pastures in the 1970s. In wheat fields, introduced varieties of
ryegrass, while good for grazing sheep, are intense competitors with wheat. Ryegrasses produce so
many seeds that, if left unchecked, they can completely choke a field. Herbicides provided excel-
lent control, while reducing soil disrupting because of less need to plough. Within little more than
a decade, ryegrass and other weeds began to develop resistance. Australian farmers evolved again
and began diversifying their techniques.

In 1983, patches of ryegrass had become immune to Hoegrass, a family of herbicides that inhibit
an enzyme called acetyl coenzyme A carboxylase.

Ryegrass populations were large, and had substantial genetic diversity, because farmers had planted
many varieties. Ryegrass is cross-pollinated by wind, so genes shuffle frequently. Farmers sprayed
inexpensive Hoegrass year after year, creating selection pressure, but were diluting the herbicide in
order to save money, increasing plants survival. Hoegrass was mostly replaced by a group of herbi-
cides that block acetolactate synthase, again helped by poor application practices. Ryegrass evolved
a kind of “cross-resistance” that allowed it to rapidly break down a variety of herbicides. Australian
farmers lost four classes of herbicides in only a few years. As of 2013 only two herbicide classes,
called Photosystem II and long-chain fatty acid inhibitors, had become the last hope.

Aquatic Weed Harvester

An aquatic weed harvester in action on a freshwater lake

An aquatic weed harvester

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Weed Control: Methods and Techniques 195

An aquatic weed harvester, also known as a water mower, mowing boat and weed cutting boat, is
an aquatic machine specifically designed for inland watercourse management to cut and harvest
underwater weeds, reeds and other aquatic plant life. The action of removing aquatic plant life in
such a manner has been referred to as “aquatic harvesting”.

Overview
Water is an important resource and in many countries, waterways are increasingly clogged by
aquatic plant growth. This is particularly so in tropical countries where warmer water means the
plants grow more quickly, and increasing run-off of fertilisers and effluent has exacerbated the
problem. Irrigation ditches and pumps can become overgrown with vegetation, power station and
factory water intakes can get blocked, boats can get hindered, fish stocks can be disrupted, and
water moves more slowly, resulting in greater evapotranspiration and a greater risk of flooding. In
some large irrigation projects in India, canals have become so overgrown with vegetation that wa-
ter flow has been reduced to a fifth of its previous amount. In Bangladesh, floodwater has washed
mats of water hyacinth onto paddy fields, overwhelming the emerging rice crops. Small fish can
become entangled in excessive algal growth.

A pond in Odessa, Ukraine covered with water lettuce (Pistia stratiotes)

Rice is the main aquatic plant grown for human food, but smaller areas of watercress and water
chestnut are also cultivated. In their native environments, aquatic weeds are part of a balanced
ecosystem, and it is mainly introduced species of water plant that become invasive and cause prob-
lems by congesting water bodies. The worst culprits, found in both temperate and tropical wa-
terways, are floating plants such as water hyacinth, water lettuce and Salvinia, fully submerged
rooting plants such as Hydrilla and water milfoil and rooting plants that reach the surface such as
cattail, papyrus, bulrush and reed.

Weed Harvesting Equipment


Weed cutting boats are developed to enable the maintenance of canals, lakes and rivers and to re-
move excessive aquatic life such as algae and other plants that may negatively affect a waterway’s
ecology. Mechanical harvesters are large floating machines that have underwater cutting blades
that sever the stems of underwater plants, gather the weeds and raise them on conveyor belts,
storing the vegetation on board in a hold. Periodically this is discharged to a barge or an onshore
facility. The harvested product can be composted, sent to a landfill site or used in land reclamation.

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196 Fundamentals of Weed Science

In developing countries aquatic vegetation may be harvested by hand or by net from the shore, cut
and harvested by boat and lifted ashore by hand, crane, pump or conveyor system. The harvested
vegetation may be used for the feeding of livestock. To reduce the high moisture content and to
make it easier to transport, the weed can be chopped and pressed. Other uses to which the harvest-
ed vegetation can be put include ensiling the material for livestock fodder, adding it to the soil as a
bulky organic fertilizer, manufacturing the raw material into pulp, paper or fibre, and fermenting
it to produce methane for energy production.

Advantages and Disadvantages


Mechanical harvesters can be effective at clearing aquatic weeds but the machines are expensive
and the process may need to be repeated several times in a growing season. Small fragments of
weed remain in the water and may spread to other locations thereby aiding in the dispersal of in-
vasive species. Some areas may be too shallow for the mechanical harvester and it may be unable
to access restricted locations. Submerged tree stumps can damage the machine. An alternative to
mechanical harvesting is the use of herbicides, which are easy to apply and less expensive, but may
have unwanted impacts on the environment.

By Region
Africa

Water hyacinth growing in Lake Victoria at Kisumu

In June 2015 in Uganda, use of the aquatic weed harvester was recommended by the country’s
Ministry of Agriculture to reduce water hyacinth growth in Lake Victoria, which has caused a scar-
city of fish in the lake. The scarcity of fish has negatively-affected the livelihood of locals who live in
the lake’s region. Additional types of machines were recommended to address the problem, which
were the hydraulic harvester, take out elevator and dredger.

India
In August 2015 in Hyderabad, India, an aquatic weed harvester was used to remove algal weeds
and trash from Hussainsagar Lake. An amphibious hydraulic excavator was also used. The work
was performed to address pollution problems at the lake, which had accumulated significant algae
and rubbish such as plastic bags and plastic waste, food wrappers, and various garbage, some of
which was floating atop the lake.

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Weed Control: Methods and Techniques 197

A mowing boat

United States
Some U.S. companies manufacture aquatic weed harvesters. One such machine manufactured by
a U.S. company can cut around one-half an acre of weeds a day, and costs over $100,000.

Tampa, Florida used an aquatic weed harvester in 2013 to clear aquatic plant life from lakes in the
area.

In 2012 in Leoni, Michigan, an aquatic weed harvester was used to remove algae from Center Lake.
The algae appeared to have roots, and other techniques such as attempting to kill it with chemicals
were unsuccessful. The machine used collects algae and weeds that live at the bottom of the lake,
removing them from the waterway.

Soil Steam Sterilization


Soil steam sterilization (soil steaming) is a farming technique that sterilizes soil with steam in open
fields or greenhouses. Pests of plant cultures such as weeds, bacteria, fungi and viruses are killed
through induced hot steam which causes their cell structure to physically degenerate. Biologically,
the method is considered a partial disinfection. Important heat-resistant, spore-forming bacteria
survive and revitalize the soil after cooling down. Soil fatigue can be cured through the release of
nutritive substances blocked within the soil. Steaming leads to a better starting position, quicker
growth and strengthened resistance against plant disease and pests. Today, the application of hot
steam is considered the best and most effective way to disinfect sick soil, potting soil and compost.
It is being used as an alternative to bromomethane, whose production and use was curtailed by
the Montreal Protocol. “Steam effectively kills pathogens by heating the soil to levels that cause
protein coagulation or enzyme inactivation.”

Benefits of Soil Steaming


Soil sterilization provides secure and quick relief of soils from substances and organisms harmful
to plants such as:
• Bacteria

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198 Fundamentals of Weed Science

• Viruses
• Fungi
• Nematodes and
• Other Pests
Further positive effects are:
• All weed and weed seeds are killed
• Significant increase of crop yields
• Relief from soil fatigue through activation of chemical – biological reactions
• Blocked nutritive substances in the soil are tapped and made available for plants
• Alternative to Methyl Bromide and other critical chemicals in agriculture

Steaming With Superheated Steam


Through modern steaming methods with superheated steam at 180–200 °C, an optimal soil dis-
infection can be achieved. Soil only absorbs a small amount of humidity. Micro organisms become
active once the soil has cooled down. This creates an optimal environment for instant tillage with
seedlings and seeds. Additionally the method of integrated steaming can promote a target-orient-
ed resettlement of steamed soil with beneficial organisms. In the process, the soil is first freed from
all organisms and then revitalized and microbiologically buffered through the injection of a soil
activator based on compost which contains a natural mixture of favorable microorganisms (e.g.
Bacillus subtilis, etc.).

Different types of such steam application are also available in practice, including substrate steam-
ing and surface steaming.

Surface Steaming
Several methods for surface steaming are in use amongst which are: area sheet steaming, the
steaming hood, the steaming harrow, the steaming plough and vacuum steaming with drainage
pipes or mobile pipe systems.

In order to pick the most suitable steaming method, certain factors have to be considered such as
soil structure, plant culture and area performance. At present, more advanced methods are being
developed, such as sandwich steaming or partially integrated sandwich steaming in order to min-
imize energy and cost as much as possible.

Sheet Steaming
Large area sheet steaming in greenhouses using a steam injector.

Surface steaming with special sheets (sheet steaming) is a method which has been established for
decades in order to steam large areas reaching from 15 to 400 m² in one step. If properly applied,
sheet steaming is simple and highly economic. The usage of heat resistant, non-decomposing in-

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Weed Control: Methods and Techniques 199

sulation fleece saves up to 50% energy, reduces the steaming time significantly and improves pen-
etration. Single working step areas up to 400 m² can be steamed in 4–5 hours down to 25–30 cm
depth / 90°C. The usage of heat resistant and non-decomposing synthetic insulation fleece, 5 mm
thick, 500 gr / m², can reduce steaming time by about 30%. Through a steam injector or a per-
forated pipe, steam is injected underneath the sheet after it has been laid out and weighted with
sand sacks.

The area performance in one working step depends on the capacity of the steam generator (e.g.
steam boiler):

Steam capacity kg/h: 100 250 300 400 550 800 1000 1350 2000

30- 50- 60-


Area m²: 15-20 80-120 130-180 180-220 220-270 300-400
50 65 90

The steaming time depends on soil structure as well as outside temperature and amounts to 1-1.5
hours per 10 cm steaming depth. Hereby the soil reaches a temperature of about 85°C. Milling
for soil loosening is not recommended since soil structure may become too fine which reduces its
penetrability for steam. The usage of spading machines is ideal for soil loosening. The best results
can be achieved if the soil is cloddy at greater depth and granulated at lesser depth.

In practice, working with at least two sheets simultaneously has proven to be highly effective.
While one sheet is used for steaming the other one is prepared for steam injection, therefore un-
necessary steaming recesses are avoided.

Depth Steaming With Vacuum


Steaming with vacuum which is induced through a mobile or fixed installed pipe system in the
depth of the area to be steamed, is the method that reaches the best penetration. Despite high
capital cost, the fixed installation of drainage systems is reasonable for intensively used areas since
steaming depths of up to 80 cm can be achieved.

In contrast to fixed installed drainage systems, pipes in mobile suction systems are on the surface.
A central suction pipeline consisting of zinc-coated, fast-coupling pipes are connected in a regular
spacing of 1.50 m and the ends of the hoses are pushed into the soil to the desired depth with a
special tool.

The steaming area is covered with a special steaming sheet and weighted all around as with sheet
steaming. The steam is injected underneath the sheet through an injector and protection tunnel.
While with short areas up to 30 m length steam is frontally injected, with longer areas steam is
induced in the middle of the beet using a T-connection branching out to both sides. As soon as
the sheet is inflated to approximately 1m by the steam pressure, the suction turbine is switched
on. First, the air in the soil is removed via the suction hoses. A vacuum is formed and the steam is
pulled downward.

During the final phase, when the required steaming depth has been reached, the ventilator runs
non-stop and surplus steam is blown out. To ensure that this surplus steam is not lost, it is fed back
under the sheet.

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200 Fundamentals of Weed Science

As with all other steaming systems, a post-steaming period of approximately 20–30 minutes is re-
quired. Steaming time is approximately 1 hour per 10 cm steaming depth. The steam requirement
is approximately 7–8 kg/m².

The most important requirement, as with all steaming systems, is that the soil is well loosened
before steaming, to ensure optimal penetration.

Negative Pressure Technique


“Negative Pressure technique generates appropriate soil temperature at a 60 cm depth and com-
plete control of nematodes, fungi and weeds is achieved. In this technique, the steam is introduced
under the steaming sheath and forced to enter the soil profile by a negative pressure. The negative
pressure is created by a fan that sucks the air out of the soil through buried perforated polypro-
pylene pipes. This system requires a permanent installation of perforated pipes into the soil, at a
depth of at least 60 cm to be protected from plough.”

Steaming with Hoods

Half automatic steaming hood with three wings in greenhouse

A steaming hood is a mobile device consisting of corrosion-resistant materials such as aluminum,


which is put down onto the area to be steamed. In contrast to sheet steaming, cost-intensive work-
ing steps such as laying out and weighting the sheets don’t occur, however the area steamed per
working step is smaller in accordance to the size of the hood.

Outdoors, a hood is positioned either manually or via tractor with a special pre-stressed 4 point
suspension arm. Steaming time amounts to 30 min for a penetration down to 25 cm depth. Here-
by a temperature of 90°C can be reached. In large stable glasshouses, the hoods are attached to
tracks. They are lifted and moved by pneumatic cylinders. Small and medium-sized hoods up to
12m² are lifted manually using a tipping lever or moved electrically with special winches.

Combined Surface and Depth Injection of Steam (Sandwich Steaming)


Sandwich steaming, which was developed in a project among DEIAFA, University of Turin (Italy,
www.deiafa.unito.it) and Ferrari Costruzioni Meccaniche, represents a combination of depth and
surface steaming, offers an efficient method to induce hot steam into the soil. The steam is simulta-
neously pushed into the soil from the surface and from the depth. For this purpose, the area, which

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Weed Control: Methods and Techniques 201

must be equipped with a deep steaming injection system, is covered with a steaming hood. The
steam enters the soil from the top and the bottom at the same time. Sheets are not suitable, since
a high pressure up to 30 mm water column arises underneath the cover.

Sandwich steaming offers several advantages. On the one hand, application of energy can be in-
creased to up to 120  kg steam per m²/h. In comparison to other steaming methods up to 30%
energy savings can be achieved and the usage of fuel (e.g. heating oil) accordingly decreases. The
increased application of energy leads to a quick heating of the soil which reduces the loss of heat.
On the other hand, only half of the regular steaming time is needed.

Sandwich steaming machine model Sterilter constructed by Ferrari Costruzioni Meccaniche equipped with MSD/
moeschle steam boiler

Comparison of sandwich steaming with other steam injection methods relating to steam output
and energy demand(*):

Steaming method max. steam output energy demand (*)

Sheet steaming 6 kg/m2h about 100 kg steam/m3

Depth steaming (Sheet + vacuum) 14 kg/m2h about 120 kg steam/m3

Hood steaming (Alu) 30 kg/m2h about 80 kg steam/m3

Hood steaming (Steel) 50 kg/m2h about 75 kg steam/m3

Sandwich steaming 120 kg/m2h about 60 kg steam/m3

(*) in soil max 30% moisture

Clearly, Sandwich steaming reaches the highest steam output at the lowest energy demand.

Partially Integrated Sandwich Steaming


The partial integrated Sandwich steaming is an advanced combined method for steaming merely
the areas which shall be planted and purposely leaving out those areas which shall not be used.
In order to avoid risk of re-infection of steamed areas with pest from unsteamed areas, beneficial
organisms can directly be injected into the hygenized soil via a soil activator (e.g. special compost).
The partial sandwich steaming unlocks further potential savings in the steaming process.

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202 Fundamentals of Weed Science

Container / Stack Steaming


Stack steaming is used when thermically treating compost and substrates such as turf. Depending
on the amount, the material to be steamed is piled up to 70 cm height in steaming boxes or in small
dump trailers. Steam is evenly injected via manifolds. For huge amounts, steaming containers and
soil boxes are used which are equipped with suction systems to improve steaming results. Midget
amounts can be steamed in special small steaming devices.

The amount of soil steamed should be tuned in a way that steaming time amounts to at most 1.5 h
in order to avoid large quantities of condensed water in the bottom layers of the soil.

Steam Output kg/h: 100 250 300 400 550 800 1000 1350 2000
1.0- 2.5- 3.0- 4.0- 5.5- 8.0- 10.0- 20.0-
m3/h about: 14.0-18.0
1.5 3.0 3.5 5.0 7.0 10.0 13.0 25.0

In light substrates, such as turf, the performance per hour is significantly higher.

History
Ancient civilizations in India and Egypt used steam, generated through the targeted usage of inci-
dent solar radiation on watered top soil, to sanitize and revive their arable land.

Modern soil steam sterilization was first discovered in 1888 (by Frank in Germany) and was first
commercially used in the United States (by Rudd) in 1893 (Baker 1962). Since then, a wide vari-
ety of steam machines have been built to disinfest both commercial greenhouse and nursery field
soils (Grossman and Liebman 1995). In the 1950s, for example, steam sterilization technologies
expanded from disinfestation of potting soil and greenhouse mixes to commercial production of
steam rakes and tractor-drawn steam blades for fumigating small acres of cut flowers and other
high-value field crops (Langedijk 1959). Today, even more effective steam technologies are being
developed.

Application of Hot Steam


• In horticulture as well as nurseries for sterilization of substrates and top soil
• In agriculture for sterilization and treatment of food waste for pig fattening and heating of
molasses
• In mushroom cultivation for pasteurization of growing rooms, sterilization of top soil and
combined application as heating
• In wineries as combination boiler for sterilization and cleaning of storage tanks, tempering
of mash and for warm water generation.

Soil Solarization
Soil solarization is an environmentally friendly method of using solar power for controlling pests

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Weed Control: Methods and Techniques 203

such as soilborne plant pathogens including fungi, bacteria, nematodes, and insect and mite pests
along with weed seed and seedlings in the soil by mulching the soil and covering it with tarp, usu-
ally with a transparent polyethylene cover, to trap solar energy. It may also describe methods of
decontaminating soil using sunlight or solar power. This energy causes physical, chemical, and
biological changes in the soil.

Soil Disinfestation
Soil solarization is a relatively new soil disinfestation method, first described in extensive scientific
detail by Katan et al. in 1976, presenting the results of a series of studies performed under field
conditions, initiated in 1973, for controlling soilborne pathogens and weeds, mostly as a pre-plant-
ing soil treatment. Soil is mulched and then covered with transparent polyethylene during the hot
season, thereby heating it and killing the pests.

Soil Decontamination
A 2008 study used a solar cell to generate an electric field for electrokinetic (EK) remediation of
cadmium-contaminated soil. The solar cell could drive the electromigration of cadmium in con-
taminated soil, and the removal efficiency that was achieved by the solar cell was comparable with
that achieved by conventional power supply.

In Korea, various remediation methods of soil slurry and groundwater contaminated with ben-
zene at a polluted gas station site were evaluated, including a solar-driven, photocatalyzed reactor
system along with various advanced oxidation processes (AOP). The most synergistic remediation
method incorporated a solar light process with TiO2 slurry and H2O2 system, achieving 98% ben-
zene degradation, a substantial increase in the removal of benzene.

History
Attempts were made to use solar energy for controlling disease agents in soil and in plant material
already in the ancient civilization of India. In 1939, Groashevoy, who used the term “solar energy
for sand disinfection,” controlled Thielaviopsis basicola upon heating the sand by exposure to di-
rect sunlight.

Soil solarization is the third approach for soil disinfestation; the two other main approaches, soil
steaming and fumigation; were developed at the end of the 19th century. The idea of solariza-
tion was based on observations by extension workers and farmers in the hot Jordan Valley, who
noticed the intensive heating of the polyethylene-mulched soil. The involvement of biological
control mechanisms in pathogen control and the possible implications were indicated in the first
publication, noticing the very long effect of the treatment. In 1977, American scientists from the
University of California at Davis reported the control of Verticillium in a cotton field, based on
studies started in 1976, thus denoting, for the first time, the possible wide applicability of this
method.

The use of polyethylene for soil solarization differs in principle from its traditional agricultural
use. With solarization, soil is mulched during the hottest months (rather than the coldest, as in

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204 Fundamentals of Weed Science

conventional plasticulture which is aimed at protecting the crop) in order to increase the maximal
temperatures in an attempt to achieve lethal heat levels.

In the first 10 years following the influential 1976 publication, soil solarization was investigated
in at least 24 countries and has been now been applied in more than 50, mostly in the hot re-
gions, although there were some important exceptions. Studies have demonstrated effectiveness
of solarization with various crops, including vegetables, field crops, ornamentals and fruit trees,
against many pathogens, weeds and a soil arthropod. Those pathogens and weeds which are not
controlled by solarization were also detected. The biological, chemical and physical changes that
take in solarized soil during and after the solarization have been investigated, as well as the inter-
action of solarization with other methods of control. Long-term effects including biological control
and increased growth response were verified in various climatic regions and soils, demonstrating
the general applicability of solarization. Computerized simulation models have been developed
to guide researchers and growers whether the ambient conditions of their locality are suitable for
solarization.

Studies of the improvement of solarization by integrating it with other methods or by solarizing


in closed glasshouses, or studies concerning commercial application by developing mulching ma-
chines were also carried out.

The use of solarization in existing orchards (e.g. controlling Verticillium in pistachio plantations)
is an important deviation from the standard preplanting method and was reported as early as 1979.

Stale Seed Bed


A false or stale seed bed is a useful weed control technique which involves creating a seedbed some
weeks before seed is due to be sown. Preparation of such seedbed makes sure that any weed seeds
that have been disturbed and brought to the soil surface during cultivation will thus have a chance
to germinate, and can then be hoed off or eliminated with the use of a flame weeder before sowing
of the actual crop is carried out.

The technique can be utilized in early spring, when the weather is still too cold for proper seed
germination. Several passes are made with a rototiller or plow, then weed seeds are allowed to
germinate as weather permits.

By tilling, the farmer increases the chance of weed seed germination by the same method as
one would for favorable vegetable/crops: the fine soil allows weed seed to grow rapidly by
allowing the seed to open and the roots to spread easier than in compacted soil. Deep tilling
will also bring dormant seed to the surface for germination. Some species of plant are known
for seeds that can lay deeply buried in the soil for years before favorable conditions allow ger-
mination.

Timing is important, however. Weed seeds must be tilled/howed or otherwise destroyed before
they themselves can create new seeds. By destroying them early, the farmer eliminates most of
that season’s annual weeds. Turning the dead weeds back into the soil also increases soil nutrient
content, although this difference is slight.

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Weed Control: Methods and Techniques 205

In many cases, several tillings are done, perhaps every two weeks beginning in very early spring.
This allows more and more weed seeds to germinate only to be killed off later. This eliminates
more weeds, but care must be used to not delay planting of a desirable crop later than the crop
needs for a successful season’s growth.

After several years, most, if not all, weeds can be eliminated from the seed bank in the soil. In some
cases the effect can be noticed in the same year the process is first carried out.

If the weed patch is vigorous enough, the farmer may mow the patch first before tilling. This allows
for easier/quicker decomposition in the soil when the plants are turned under.

Some farmers are noted for applying a light and inexpensive fertilizer mix to the soil to cause even
more weed seeds to germinate. Although the merits of this may seem a trifle at first, it can pay
dividends in the long run by eliminating seeds earlier that otherwise would have sprouted in later
years. This too is open to debate.

Drip Irrigation

An Emitter or dripper in action

Open pressure compensated dripper

Drip irrigation is a form of irrigation that saves water and fertilizer by allowing water to drip slow-
ly to the roots of many different plants, either onto the soil surface or directly onto the root zone,
through a network of valves, pipes, tubing, and emitters. It is done through narrow tubes that
deliver water directly to the base of the plant. It is chosen instead of surface irrigation for various
reasons, often including concern about minimizing evaporation.

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206 Fundamentals of Weed Science

History
Primitive drip irrigation has been used since ancient times. Fan Sheng-Chih Shu, written in China
during the first century BCE, describes the use of buried, unglazed clay pots filled with water as
a means of irrigation. Modern drip irrigation began its development in Germany in 1860 when
researchers began experimenting with subsurface irrigation using clay pipe to create combination
irrigation and drainage systems. Research was later expanded in the 1920s to include the applica-
tion of perforated pipe systems. The usage of plastic to hold and distribute water in drip irrigation
was later developed in Australia by Hannis Thill.

Drip irrigation in Mexico vineyard, 2000

Usage of a plastic emitter in drip irrigation was developed in Israel by Simcha Blass and his son
Yeshayahu. Instead of releasing water through tiny holes easily blocked by tiny particles, water
was released through larger and longer passageways by using velocity to slow water inside a plastic
emitter. The first experimental system of this type was established in 1959 by Blass who partnered
later (1964) with Kibbutz Hatzerim to create an irrigation company called Netafim. Together they
developed and patented the first practical surface drip irrigation emitter.

In the United States, the first drip tape, called Dew Hose, was developed by Richard Chapin of
Chapin Watermatics in the early 1960s.

Modern drip irrigation has arguably become the world’s most valued innovation in agriculture
since the invention of the impact sprinkler in the 1930s, which offered the first practical alternative
to surface irrigation. Drip irrigation may also use devices called micro-spray heads, which spray
water in a small area, instead of dripping emitters. These are generally used on tree and vine crops
with wider root zones. Subsurface drip irrigation (SDI) uses permanently or temporarily buried
dripperline or drip tape located at or below the plant roots. It is becoming popular for row crop irri-
gation, especially in areas where water supplies are limited or recycled water is used for irrigation.
Careful study of all the relevant factors like land topography, soil, water, crop and agro-climatic
conditions are needed to determine the most suitable drip irrigation system and components to be
used in a specific installation.

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Weed Control: Methods and Techniques 207

Components and Operation

Drip irrigation system layout and its parts

Water distribution in subsurface drip irrigation

Nursery flowers watered with drip irrigation in Israel

Components used in drip irrigation (listed in order from water source) include:

• Pump or pressurized water source

• Water filter(s) or filtration systems: sand separator, Fertigation systems (Venturi injector)
and chemigation equipment (optional)

• Backwash controller (Backflow prevention device)

• Pressure Control Valve (pressure regulator)

• Distribution lines (main larger diameter pipe, maybe secondary smaller, pipe fittings)

• Hand-operated, electronic, or hydraulic control valves and safety valves

• Smaller diameter polytube (often referred to as “laterals”)

• Poly fittings and accessories (to make connections)

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208 Fundamentals of Weed Science

• Emitting devices at plants (emitter or dripper, micro spray head, inline dripper or inline
driptube)

In drip irrigation systems, pump and valves may be manually or automatically operated by a con-
troller.

Most large drip irrigation systems employ some type of filter to prevent clogging of the small emit-
ter flow path by small waterborne particles. New technologies are now being offered that minimize
clogging. Some residential systems are installed without additional filters since potable water is
already filtered at the water treatment plant. Virtually all drip irrigation equipment manufacturers
recommend that filters be employed and generally will not honor warranties unless this is done.
Last line filters just before the final delivery pipe are strongly recommended in addition to any
other filtration system due to fine particle settlement and accidental insertion of particles in the
intermediate lines.

Drip and subsurface drip irrigation is used almost exclusively when using recycled municipal waste
water. Regulations typically do not permit spraying water through the air that has not been fully
treated to potable water standards.

Because of the way the water is applied in a drip system, traditional surface applications of
timed-release fertilizer are sometimes ineffective, so drip systems often mix liquid fertilizer with
the irrigation water. This is called fertigation; fertigation and chemigation (application of pesti-
cides and other chemicals to periodically clean out the system, such as chlorine or sulfuric acid)
use chemical injectors such as diaphragm pumps, piston pumps, or aspirators. The chemicals may
be added constantly whenever the system is irrigating or at intervals. Fertilizer savings of up to
95% are being reported from recent university field tests using drip fertigation and slow water de-
livery as compared to timed-release and irrigation by micro spray heads.

Properly designed, installed, and managed, drip irrigation may help achieve water conservation
by reducing evaporation and deep drainage when compared to other types of irrigation such
as flood or overhead sprinklers since water can be more precisely applied to the plant roots. In
addition, drip can eliminate many diseases that are spread through water contact with the foli-
age. Finally, in regions where water supplies are severely limited, there may be no actual water
savings, but rather simply an increase in production while using the same amount of water as
before. In very arid regions or on sandy soils, the preferred method is to apply the irrigation
water as slowly as possible.

Pulsed irrigation is sometimes used to decrease the amount of water delivered to the plant at any
one time, thus reducing runoff or deep percolation. Pulsed systems are typically expensive and
require extensive maintenance. Therefore, the latest efforts by emitter manufacturers are focused
toward developing new technologies that deliver irrigation water at ultra-low flow rates, i.e. less
than 1.0 liter per hour. Slow and even delivery further improves water use efficiency without incur-
ring the expense and complexity of pulsed delivery equipment.

An emitting pipe is a type of drip irrigation tubing with emitters pre-installed at the factory with
specific distance and flow per hour as per crop distance.

An emitter restricts water flow passage through it, thus creating head loss required (to the extent

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Weed Control: Methods and Techniques 209

of atmospheric pressure) in order to emit water in the form of droplets. This head loss is achieved
by friction / turbulence within the emitter.

Advantages and Disadvantages

Drip irrigation and spare drip irrigation tubes in banana farm at Chinawal, India

Pot irrigation by On-line drippers

Pressure compensated integral dripper on soilless without growing channels

The advantages of drip irrigation are:

• Fertilizer and nutrient loss is minimized due to localized application and reduced
leaching.

• Water application efficiency is high if managed correctly

• Field levelling is not necessary.

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210 Fundamentals of Weed Science

• Fields with irregular shapes are easily accommodated.

• Recycled non-potable water can be safely used.

• Moisture within the root zone can be maintained at field capacity.

• Soil type plays less important role in frequency of irrigation.

• Soil erosion is lessened.

• Weed growth is lessened.

• Water distribution is highly uniform, controlled by output of each nozzle.

• Labour cost is less than other irrigation methods.

• Variation in supply can be regulated by regulating the valves and drippers.

• Fertigation can easily be included with minimal waste of fertilizers.

• Foliage remains dry, reducing the risk of disease.

• Usually operated at lower pressure than other types of pressurised irrigation, reducing
energy costs.

The disadvantages of drip irrigation are:

• Initial cost can be more than overhead systems.

• The sun can affect the tubes used for drip irrigation, shortening their usable life.

• If the water is not properly filtered and the equipment not properly maintained, it can re-
sult in clogging.

• For subsurface drip the irrigator cannot see the water that is applied. This may lead to the
farmer either applying too much water (low efficiency) or an insufficient amount of water,
this is particularly common for those with less experience with drip irrigation.

• Drip irrigation might be unsatisfactory if herbicides or top dressed fertilizers need sprin-
kler irrigation for activation.

• Drip tape causes extra cleanup costs after harvest. Users need to plan for drip tape winding,
disposal, recycling or reuse.

• Waste of water, time and harvest, if not installed properly. These systems require careful
study of all the relevant factors like land topography, soil, water, crop and agro-climatic
conditions, and suitability of drip irrigation system and its components.

• In lighter soils subsurface drip may be unable to wet the soil surface for germination. Re-
quires careful consideration of the installation depth.

• most drip systems are designed for high efficiency, meaning little or no leaching fraction.
Without sufficient leaching, salts applied with the irrigation water may build up in the root

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Weed Control: Methods and Techniques 211

zone, usually at the edge of the wetting pattern. On the other hand, drip irrigation avoids
the high capillary potential of traditional surface-applied irrigation, which can draw salt
deposits up from deposits below.

• the PVC pipes often suffer from rodent damage, requiring replacement of the entire tube
and increasing expenses.

• Drip irrigation systems cannot be used for damage control by night frosts (like in the case
of sprinkler irrigation systems)

Uses
Drip irrigation is used in farms, commercial greenhouses, and residential gardeners. Drip irriga-
tion is adopted extensively in areas of acute water scarcity and especially for crops and trees such
as coconuts, containerized landscape trees, grapes, bananas, pandey, eggplant, citrus, strawber-
ries, sugarcane, cotton, maize, and potatoes.

Irrigation dripper

Drip irrigation for garden available in drip kits are increasingly popular for the homeowner and
consist of a timer, hose and emitter. Hoses that are 4 mm in diameter are used to irrigate flower
pots.

Bioherbicide
A bioherbicide is a biologically based control agent for weeds. Among the three major types of
pesticides (agricultural pest-control agents) herbicides are used to control weeds, or undesirable
plants. (The other major classes of pesticides are insecticides, which control insects; and fungi-
cides, which control fungi. Normally, any pesticide can only be expected to control, but not eradi-
cate, agricultural pests.)

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212 Fundamentals of Weed Science

Almost every agricultural pest has at least one naturally occurring enemy that will reduce its pop-
ulation. Bioherbicides are weed-control agents that use such naturally occurring enemies, rather
than depending on man-made chemicals. This can be important because agents of biological con-
trol ordinarily have many fewer, and much milder, effects on the environment than do synthetic
chemicals. What is more, they tend not to lead to the public health problems that chemicals are
associated with. These two advantages of biological control agents, including bioherbicides, do not
however mean that they need not also be subjected to careful tests for environmental and public
health safety.

Bioherbicides may be compounds derived from microbes such as fungi, bacteria or protozoa; or
phytotoxic plant residues, extracts or single compounds derived from other plant species.

A bioherbicide based on a fungus is called a mycoherbicide.

In the industry, bioherbicides and other biopesticides are often referred to as “naturals”.

Bush Regeneration
Bush regeneration, a form of natural area restoration, is the term used in Australia for the ecolog-
ical restoration of remnant vegetation areas, such as through the minimisation of negative distur-
bances, both exogenous such as exotic weeds and endogenous such as erosion. It may also attempt
to recreate conditions of pre-European arrival, for example by simulating endogenous disturbanc-
es such as fire. Bush regeneration attempts to protect and enhance the floral biodiversity in an area
by providing conditions conducive to the recruitment and survival of native plants. Bushcare’s
Major Day Out is an Australian national day of community participation in the care of bushland.
In 2012 nearly 100 bushcare sites participated in this annual event. For more information go to
www.bushcaresmajordayout.org.

History
Bradley Method
In the early 1960s Joan and Eileen Bradley developed a series of weed control techniques through
a process of trial and error. Their work was the beginning of minimal disturbance bush regenera-
tion in New South Wales. The Bradley method urges a naturalistic approach by encouraging the
native vegetation to self-reestablish. The Bradleys used their method to successfully clear weeds
from a 16 hectares (40 acres) reserve in Ashton Park, part of Sydney Harbour National Park,
NSW. The process demonstrated that, following a period of consecutive ‘follow up’ treatments
of diminishing time requirement, subsequent maintenance was needed only once or twice a
year, mainly in vulnerable spots such as creek banks, roadsides, and clearings, to be maintained
weed-free.

The aim of their work was to clear small niches adjacent to healthy native vegetation such that the
each area will regenerate from in-situ soil seed banks or be re-colonised and stabilized by the re-
generation of native plants, replacing an area previously occupied by weeds. The Bradley method
follows three main principles,

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Weed Control: Methods and Techniques 213

1. secure the best areas first

2. minimise disturbance to the natural conditions (e.g. minimise soil disturbance and off-tar-
get damage).

3. don’t overclear, let the regenerative ability of the bush set the pace of clearance (Bradley
1988).

The priority securing of the best quality vegetation aids in preserving areas of top biodiversity
which provide regeneration potential to expand these areas and reclaim areas as bushland.

Modern Practice
The adoption of minimal disturbance bush regeneration increased in the decades that followed the
work of the Bradleys. Their principles have guided bushcare programs in Australia, although the in-
clusion of herbicide in modern bush regeneration is a notable deviation from the ideals of the Bradley
sisters. In addition, rather than ‘minimal disturbance’, a more favoured and ecologically sound trend
since the 1990s has been towards more ‘appropriate disturbance’ as many Australian plant commu-
nities require some level of perturbation to trigger germination from long-buried seed banks. This
has led to a range of additional disturbance-based techniques (such as burns and soil disturbance)
being included in the regenerator’s ‘tool kit’ in dry forest and grassland areas. Field experience has
found that, even in rainforest areas, a resilience to disturbance is evident, enabling regenerators to
clear weed in a fairly extensive manner to trigger rainforest recovery. This is borne out by a thriving
rainforest regeneration industry in northern NSW Australia, modelled on the pioneering work of
John Stockard at Wingham Brush (Stockard 1991, Stockard 1999). The rule of thumb in all cases is
to constrain clearing to that area that matches the project’s follow up resources.

The increased awareness and consideration of Australia’s biodiversity by citizens has incre-
mentally increased pressure on local governments to adopt conservation programs for rem-
nant vegetation on council land. Most peri urban councils now have some involvement in
bush regeneration, either through planning, land management, volunteer support or through
employment of bush regeneration practitioners. In NSW the level of coordination of bush
regeneration programs through local governments is high, although in some other areas at
present a lack of coordination is a serious concern in bush regeneration on public land, with
only 40% of councils liaising with other councils. In such areas there may be a need for stra-
tegic management at a regional scale through Natural Resource Management Boards or non
government organisations such as Trees For Life, which are involved in bushcare programs
across wider areas.

Purposes
The aim of bush regeneration, also known as ‘natural area restoration’, is to restore and main-
tain ecosystem health by facilitating the natural regeneration of indigenous flora, this is usually
achieved by selectively reducing the competitive interaction with invasive species, or mitigation of
negative influences such as weeds or erosion.

Invasive plant species are often the greatest threat to remnant vegetation, and therefore bush re-
generation is closely associated with weed abatement activities. Weed management as one aim

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214 Fundamentals of Weed Science

of bush regeneration, is used to increase native plant recruitment. The management of factors
such as fire and herbivory can be just as important, depending on the ecosystem being restored.
In recent years research and on-ground management has begun to recognize the importance of
ecosystem processes rather ecosystem composition and structure and research into other ways of
facilitating native plant recruitment is increasing.

Technique
The original Bradley method of bush regeneration focuses on facilitating native plant recruitment
from the seedbank, rather than planting seedlings or sowing seeds, as follows:

“Weeding a little at a time from the bush towards the weeds takes the pressure off the natives
under favourable conditions. Native seeds and spores are ready in the ground and the nat-
ural environment favours plants that have evolved in it. The balance is tipped back towards
regeneration. Keep it that way, by always working where the strongest area of bush meets the
weakest weeds”

Currently the term ‘bush regeneration’ includes activities other than weed removal, such as re-
planting and introducing species into an area where soil, water, or fire regimes have shifted the
type of plant appropriate to the area (e.g. a stormwater drain).

Weed species can be important habitat for native fauna (e.g. blackberry is important habitat for
wrens and the southern brown bandicoot) and this should be taken into consideration with bush
regeneration, for example by not clearing invasive species until adequate habitat alternatives have
been established nearby with native vegetation.

Problems can occur when insufficient follow-up is conducted as the success of bush regeneration
is dependent on allowing the native vegetation to regenerate in the area where weeds have been
removed.

List of Bushcare Groups


Organisations Offering Community Training in Bush Regeneration
• Trees For Life

• Campbelltown Council, NSW, Streamcare Group

• San Diego Chapter, California Native Plant Society

Reserves where Volunteer Groups Undertake Bush Regeneration


• George Kendall Riverside Park

• Whites Creek (Annandale)

• Puckeys Estate Reserve

• Mermaid Pool, Manly Vale Sydney

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Weed Control: Methods and Techniques 215

Cultivator

F210 Honda tiller

1949 Farmall C with C-254-A two-row cultivator

A cultivator pulled by a tractor in Canada in 1943

A cultivator is any of several types of farm implement used for secondary tillage. One sense of the
name refers to frames with the teeth (also called shanks) that pierce the soil as they are dragged
through it linearly. Another sense refers to machines that use rotary motion of disks or teeth to
accomplish a similar result. The rotary tiller is a principle example.

Cultivators stir and pulverize the soil, either before planting (to aerate the soil and prepare a
smooth, loose seedbed) or after the crop has begun growing (to kill weeds—controlled disturbance
of the topsoil close to the crop plants kills the surrounding weeds by uprooting them, burying their
leaves to disrupt their photosynthesis, or a combination of both). Unlike a harrow, which disturbs
the entire surface of the soil, cultivators are designed to disturb the soil in careful patterns, sparing
the crop plants but disrupting the weeds.

Cultivators of the toothed type are often similar in form to chisel plows, but their goals are differ-
ent. Cultivator teeth work near the surface, usually for weed control, whereas chisel plow shanks

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216 Fundamentals of Weed Science

work deep beneath the surface, breaking up hardpan. Consequently, cultivating also takes much
less power per shank than does chisel plowing.

Small toothed cultivators pushed or pulled by a single person are used as garden tools for small-
scale gardening, such as for the household’s own use or for small market gardens. Similarly sized
rotary tillers combine the functions of harrow and cultivator into one multipurpose machine.

Cultivators are usually either self-propelled or drawn as an attachment behind either a two-wheel
tractor or four-wheel tractor. For two-wheel tractors they are usually rigidly fixed and powered via
couplings to the tractors’ transmission. For four-wheel tractors they are usually attached by means
of a three-point hitch and driven by a power take-off (PTO). Drawbar hookup is also still common-
ly used worldwide. Draft-animal power is sometimes still used today, being somewhat common in
developing nations although rare in more industrialized economies.

History
The basic idea of soil scratching for weed control is ancient and was done with hoes or mattocks
for millennia before cultivators were developed. Cultivators were originally drawn by draft animals
(such as horses, mules, or oxen) or were pushed or drawn by people. In modern commercial agri-
culture, the amount of cultivating done for weed control has been greatly reduced via use of her-
bicides instead. However, herbicides are not always desirable—for example, in organic farming.

The powered rotary hoe was invented by Arthur Clifford Howard who, in 1912, began experiment-
ing with rotary tillage on his father’s farm at Gilgandra, New South Wales, Australia. Initially using
his father’s steam tractor engine as a power source, he found that ground could be mechanically
tilled without soil-packing occurring, as was the case with normal ploughing. His earliest designs
threw the tilled soil sideways, until he improved his invention by designing an L-shaped blade
mounted on widely spaced flanges fixed to a small-diameter rotor. With fellow apprentice Everard
McCleary, he established a company to make his machine, but plans were interrupted by World
War I. In 1919 Howard returned to Australia and resumed his design work, patenting a design with
5 rotary hoe cultivator blades and an internal combustion engine in 1920.

In March 1922, Howard formed the company Austral Auto Cultivators Pty Ltd, which later became
known as Howard Auto Cultivators. It was based in Northmead, a suburb of Sydney, from 1927.

Meanwhile, in North America during the 1910s, tractors were evolving away from traction en-
gine-sized monsters toward smaller, lighter, more affordable machines. The Fordson tractor espe-
cially had made tractors affordable and practical for small and medium family farms for the first
time in history. Cultivating was somewhat of an afterthought in the Fordson’s design, which reflected
the fact that even just bringing practical motorized tractive power alone to this market segment was
in itself a milestone. This left an opportunity for others to pursue better motorized cultivating. Be-
tween 1915 and 1920, various inventors and farm implement companies experimented with a class
of machines referred to as motor cultivators, which were simply modified horse-drawn shank-type
cultivators with motors added for self-propulsion. This class of machines found limited market suc-
cess. But by 1921 International Harvester had combined motorized cultivating with the other tasks
of tractors (tractive power and belt work) to create the Farmall, the general-purpose tractor tailored
to cultivating that basically invented the category of row-crop tractors.

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Weed Control: Methods and Techniques 217

In Australia, by the 1930s, Howard was finding it increasingly difficult to meet a growing worldwide
demand for exports of his machines. He travelled to the United Kingdom, founding the company
Rotary Hoes Ltd in East Horndon, Essex, in July 1938. Branches of this new company subsequent-
ly opened in the United States of America, South Africa, Germany, France, Italy, Spain, Brazil,
Malaysia, Australia and New Zealand. It later became the holding company for Howard Rotavator
Co. Ltd. The Howard Group of companies was acquired by the Danish Thrige Agro Group in 1985,
and in December 2000 the Howard Group became a member of Kongskilde Industries of Soroe,
Denmark.

When herbicidal weed control was first widely commercialized in the 1950s and 1960s, it played
into that era’s optimistic worldview in which sciences such as chemistry would usher in a new age
of modernity that would leave old-fashioned practices (such as weed control via cultivators) in
the dustbin of history. Thus herbicidal weed control was adopted very widely, and in some cases
too heavily and hastily. In subsequent decades, people overcame this initial imbalance and came
to realize that herbicidal weed control has limitations and externalities, and it must be managed
intelligently. It is still widely used, and probably will continue to be indispensable to affordable
food production worldwide for the foreseeable future; but its wise management includes seeking
alternate methods, such as the traditional standby of mechanical cultivation, where practical.

Industrial Use
To the extent that cultivating is done commercially today (such as in truck farming), it is usually
powered by tractors, especially row-crop tractors. Industrial cultivators can vary greatly in size and
shape, from 10 feet (3 m) to 80 feet (24 m) wide. Many are equipped with hydraulic wings that fold
up to make road travel easier and safer. Different types are used for preparation of fields before
planting, and for the control of weeds between row crops. The cultivator may be an implement
trailed after the tractor via a drawbar; mounted on the three-point hitch; or mounted on a frame
beneath the tractor. Active cultivator implements are driven by a power take-off shaft. While most
cultivator are considered a secondary tillage implement, active cultivators are commonly used
for primary tillage in lighter soils instead of plowing. The largest versions available are about 6 m
(20 ft) wide, and require a tractor with an excess of 150 horsepower (110 kW) (PTO) to drive them.

Field cultivators are used to complete tillage operations in many types of arable crop fields. The
main function of the field cultivator is to prepare a proper seedbed for the crop to be planted into,
to bury crop residue in the soil (helping to warm the soil before planting), to control weeds, and to
mix and incorporate the soil to ensure the growing crop has enough water and nutrients to grow
well during the growing season. The implement has many shanks mounted on the underside of a
metal frame, and small narrow rods at the rear of the machine that smooth out the soil surface for
easier travel later when planting. In most field cultivators, one-to-many hydraulic cylinders raise
and lower the implement and control its depth.

Row Crop Cultivators


The main function of the row crop cultivator is weed control between the rows of an established
crop. Row crop cultivators are usually raised and lowered by a three-point hitch and the depth is
controlled by gauge wheels.

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218 Fundamentals of Weed Science

Home made sweep. Notice the inner and outer “sweep” blades.

Sometimes referred to as sweep cultivators, these commonly have two center blades that cut weeds
from the roots near the base of the crop and turn over soil, while two rear sweeps further outward
than the center blades deal with the center of the row, and can be anywhere from 1 to 36 rows wide.

Garden Cultivators
Small tilling equipment, used in small gardens such as household gardens and small commercial
gardens, can provide both primary and secondary tillage. For example, a rotary tiller does both
the “plowing” and the “harrowing”, preparing a smooth, loose seedbed. It does not provide the
row-wise weed control that cultivator teeth would. For that task, there are single-person-pushable
toothed cultivators.

Variants and Trademarks

A Japanese two-wheel tractor

Rotary tillers are a type of cultivators. Rotary tillers are popular with home gardeners who want
large vegetable gardens. The garden may be tilled a few times before planting each crop. Rotary
tillers may be rented from tool rental centers for single-use applications, such as when planting
grass.

A small rotary hoe for domestic gardens was known by the trademark Rototiller and another,
made by the Howard Group, who produced a range of rotary tillers, was known as the Rotavator.

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Weed Control: Methods and Techniques 219

Rototiller

The small rototiller is typically propelled forward via a (1–5 horsepower or 0.8–3.5 kilo-
watts) petrol engine rotating the tines, and do not have powered wheels, though they may
have small transport/level control wheel(s). To keep the machine from moving forward too
fast, an adjustable tine is usually fixed just behind the blades so that through friction with
deeper un-tilled soil, it acts as a brake, slowing the machine and allowing it to pulverize
the soils. The slower a rototiller moves forward, the more soil tilth can be obtained. The
operator can control the amount of friction/braking action by raising and lowering the
handlebars of the tiller. Rototillers do not have a reverse as such backwards movement to-
wards the operator could cause serious injury. While operating, the rototiller can be pulled
backwards to go over areas that were not pulverized enough, but care must be taken to en-
sure that the operator does not stumble and pull the rototiller on top of himself. Rototilling
is much faster than manual tilling, but notoriously difficult to handle and exhausting work,
especially in the heavier and higher horsepower models. If the rototiller’s blades catch on
unseen subsurface objects, such as tree roots and buried garbage, it can cause the rototiller
to abruptly and violently move in any direction.

Rotavator

Unlike the Rototiller, the self-propelled Howard Rotavator is equipped with a gearbox and
driven forward, or held back, by its wheels. The gearbox enables the forward speed to be ad-
justed while the rotational speed of the tines remains constant which enables the operator to
easily regulate the extent to which soil is engaged. For a two-wheel tractor rotavator this greatly
reduces the workload of the operator as compared to a rototiller. These rotavators are generally
more heavy duty, come in higher power (4–18 horsepower or 3–13 kilowatts) with either petrol
or diesel engines and can cover much more area per hour. The trademarked word “Rotavator”
is one of the longest single-word palindromes in the English language.

Mini tiller

Mini tillers are a new type of small agricultural tillers or cultivators used by farmers or
homeowners. These are also known as power tillers or garden tillers. Compact, powerful
and, most importantly, inexpensive, these agricultural rotary tillers are providing alter-
natives to four-wheel tractors and in the small farmers’ fields in developing countries are
more economical than four-wheel tractors.

Two-wheel tractor

The higher power “riding” rotavators cross out of the home garden category into farming
category, especially in Asia, Africa and South America, capable of preparing 1 hectare of
land in 8–10 hours. These are also known as power tillers or walking tractors. Years ago
they were considered only useful for rice growing areas, where they were fitted with steel
cage-wheels for traction, but now the same are being used in both wetland and dryland
farming all over the world. They have multiple functions with related tools for dryland or
paddys, pumping, transportation, threshing, ditching, spraying pesticide. They can be used
on hills, mountains, in greenhouses and orchards. Diesel designs are more popular in de-
veloping countries than gasoline.

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220 Fundamentals of Weed Science

Mulch
A mulch is a layer of material applied to the surface of an area of soil. Its purpose is any or all of
the following:

• to conserve moisture

• to improve the fertility and health of the soil

• to reduce weed growth

• to enhance the visual appeal of the area

A mulch is usually but not exclusively organic in nature. It may be permanent (e.g. plastic sheeting)
or temporary (e.g. bark chips). It may be applied to bare soil, or around existing plants. Mulches of
manure or compost will be incorporated naturally into the soil by the activity of worms and other
organisms. The process is used both in commercial crop production and in gardening, and when
applied correctly can dramatically improve soil productivity.

Uses
Many materials are used as mulches, which are used to retain soil moisture, regulate soil tempera-
ture, suppress weed growth, and for aesthetics. They are applied to the soil surface, around trees,
paths, flower beds, to prevent soil erosion on slopes, and in production areas for flower and vege-
table crops. Mulch layers are normally two inches or more deep when applied.

They are applied at various times of the year depending on the purpose. Towards the beginning
of the growing season mulches serve initially to warm the soil by helping it retain heat which is
lost during the night. This allows early seeding and transplanting of certain crops, and encourages
faster growth. As the season progresses, mulch stabilizes the soil temperature and moisture, and
prevents the growing of weeds from seeds. In temperate climates, the effect of mulch is dependent
upon the time of year they are applied and when applied in fall and winter, are used to delay the
growth of perennial plants in the spring or prevent growth in winter during warm spells, which
limits freeze thaw damage.

The effect of mulch upon soil moisture content is complex. Mulch forms a layer between the soil
and the atmosphere which prevents sunlight from reaching the soil surface, thus reducing evap-
oration. However, mulch can also prevent water from reaching the soil by absorbing or blocking
water from light rains.

In order to maximise the benefits of mulch, while minimizing its negative influences, it is often ap-
plied in late spring/early summer when soil temperatures have risen sufficiently, but soil moisture
content is still relatively high. However, permanent mulch is also widely used and valued for its
simplicity, as popularized by author Ruth Stout, who said, “My way is simply to keep a thick mulch
of any vegetable matter that rots on both sides of my vegetable and flower garden all year long. As
it decays and enriches the soils, I add more.”

Plastic mulch used in large-scale commercial production is laid down with a tractor-drawn or

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Weed Control: Methods and Techniques 221

standalone layer of plastic mulch. This is usually part of a sophisticated mechanical process, where
raised beds are formed, plastic is rolled out on top, and seedlings are transplanted through it. Drip
irrigation is often required, with drip tape laid under the plastic, as plastic mulch is impermeable
to water.

Materials

Rubber mulch nuggets in a playground. The white fibers are nylon cords, which are present in the tires
from which the mulch is made.

Shredded wood used as mulch. This type of mulch is often dyed to improve its appearance in the landscape.

Spring daffodils push through shredded wood mulch

Materials used as mulches vary and depend on a number of factors. Use takes into consideration
availability, cost, appearance, the effect it has on the soil—including chemical reactions and pH,
durability, combustibility, rate of decomposition, how clean it is—some can contain weed seeds or
plant pathogens.

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222 Fundamentals of Weed Science

A variety of materials are used as mulch:

• Organic residues: grass clippings, leaves, hay, straw, kitchen scraps comfrey, shredded
bark, whole bark nuggets, sawdust, shells, woodchips, shredded newspaper, cardboard,
wool, animal manure, etc. Many of these materials also act as a direct composting system,
such as the mulched clippings of a mulching lawn mower, or other organics applied as
sheet composting.

• Compost: fully composted materials are used to avoid possible phytotoxicity problems.
Materials that are free of seeds are ideally used, to prevent weeds being introduced by the
mulch.

• Old carpet (synthetic or natural): makes a free, readily available mulch.

• Rubber mulch: made from recycled tire rubber.

• Plastic mulch: crops grow through slits or holes in thin plastic sheeting. This method is pre-
dominant in large-scale vegetable growing, with millions of acres cultivated under plastic
mulch worldwide each year (disposal of plastic mulch is cited as an environmental prob-
lem).

• Rock and gravel can also be used as a mulch. In cooler climates the heat retained by rocks
may extend the growing season.

In some areas of the United States, such as central Pennsylvania and northern California, mulch
is often referred to as “tanbark”, even by manufacturers and distributors. In these areas, the word
“mulch” is used specifically to refer to very fine tanbark or peat moss.

Organic Mulches
Organic mulches decay over time and are temporary. The way a particular organic mulch decom-
poses and reacts to wetting by rain and dew affects its usefulness.

Some mulches such as straw, peat, sawdust and other wood products may for a while negatively
affect plant growth because of their wide carbon to nitrogen ratio, because bacteria and fungi that
decompose the materials remove nitrogen from the surrounding soil for growth. However, whether
this effect has any practical impact on gardens is disputed by researchers and the experience of gar-
deners. Organic mulches can mat down, forming a barrier that blocks water and air flow between the
soil and the atmosphere. Vertically applied organic mulches can wick water from the soil to the sur-
face, which can dry out the soil. Mulch made with wood can contain or feed termites, so care must be
taken about not placing mulch too close to houses or building that can be damaged by those insects.
Some mulch manufacturers recommend putting mulch several inches away from buildings.

Commonly available organic mulches include:

Leaves
• Leaves from deciduous trees, which drop their foliage in the autumn/fall. They tend to be
dry and blow around in the wind, so are often chopped or shredded before application. As

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Weed Control: Methods and Techniques 223

they decompose they adhere to each other but also allow water and moisture to seep down
to the soil surface. Thick layers of entire leaves, especially of maples and oaks, can form a
soggy mat in winter and spring which can impede the new growth lawn grass and other
plants. Dry leaves are used as winter mulches to protect plants from freezing and thawing
in areas with cold winters, they are normally removed during spring.

Grass Clippings
• Grass clippings, from mowed lawns are sometimes collected and used elsewhere as mulch.
Grass clippings are dense and tend to mat down, so are mixed with tree leaves or rough
compost to provide aeration and to facilitate their decomposition without smelly putre-
faction. Rotting fresh grass clippings can damage plants; their rotting often produces a
damaging buildup of trapped heat. Grass clippings are often dried thoroughly before appli-
cation, which mediates against rapid decomposition and excessive heat generation. Fresh
green grass clippings are relatively high in nitrate content, and when used as a mulch,
much of the nitrate is returned to the soil, but the routine removal of grass clippings from
the lawn results in nitrogen deficiency for the lawn.

Peat Moss
• Peat moss, or sphagnum peat, is long lasting and packaged, making it convenient and pop-
ular as a mulch. When wetted and dried, it can form a dense crust that does not allow water
to soak in. When dry it can also burn, producing a smoldering fire. It is sometimes mixed
with pine needles to produce a mulch that is friable. It can also lower the pH of the soil
surface, making it useful as a mulch under acid loving plants.

However peat bogs are a valuable wildlife habitat, and peat is also one of the largest stores of car-
bon (in Britain, out of a total estimated 9952 million tonnes of carbon in British vegetation and
soils, 6948 million tonnes carbon are estimated to be in Scottish, mostly peatland, soils), so gar-
deners who wish to protect the environment will choose more sustainable alternatives.

Wood Chips
• Wood chips are a byproduct of the pruning of trees by arborists, utilities and parks; they
are used to dispose of bulky waste. Tree branches and large stems are rather coarse after
chipping and tend to be used as a mulch at least three inches thick. The chips are used to
conserve soil moisture, moderate soil temperature and suppress weed growth. The decay
of freshly produced chips from recently living woody plants, consumes nitrate; this is often
off set with a light application of a high-nitrate fertilizer. Wood chips are most often used
under trees and shrubs. When used around soft stemmed plants, an unmulched zone is left
around the plant stems to prevent stem rot or other possible diseases. They are often used
to mulch trails, because they are readily produced with little additional cost outside of the
normal disposal cost of tree maintenance. Wood chips come in various colors.

Woodchip Mulch
• Woodchip mulch is a byproduct of reprocessing used (untreated) timber (usually packag-

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224 Fundamentals of Weed Science

ing pallets), to dispose of wood waste by creating woodchip mulch. The chips are used to
conserve soil moisture, moderate soil temperature and suppress weed growth. Woodchip
mulch is often used under trees, shrubs or large planting areas and can last much longer
than arborist mulch. In addition, many consider woodchip mulch to be visually appealing,
as it comes in various colors. Woodchips can also be reprocessed into playground wood-
chip to be used as an impact-attenuating playground surfacing.

Bark Chips

Bark chips

• Bark chips of various grades are produced from the outer corky bark layer of timber trees.
Sizes vary from thin shredded strands to large coarse blocks. The finer types are very at-
tractive but have a large exposed surface area that leads to quicker decay. Layers two or
three inches deep are usually used, bark is relativity inert and its decay does not demand
soil nitrates. Bark chips are also available in various colors.

Straw Mulch / Field Hay / Salt Hay

Permaculture garden with a fruit tree, herbs, flowers and vegetables mulched with hay

• Straw mulch or field hay or salt hay are lightweight and normally sold in compressed
bales. They have an unkempt look and are used in vegetable gardens and as a winter cov-
ering. They are biodegradable and neutral in pH. They have good moisture retention and
weed controlling properties but also are more likely to be contaminated with weed seeds.
Salt hay is less likely to have weed seeds than field hay. Straw mulch is also available in
various colors.

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Weed Control: Methods and Techniques 225

Cardboard / Newspaper
• Cardboard or newspaper can be used as mulches. These are best used as a base layer
upon which a heavier mulch such as compost is placed to prevent the lighter cardboard/
newspaper layer from blowing away. By incorporating a layer of cardboard/newspaper
into a mulch, the quantity of heavier mulch can be reduced, whilst improving the weed
suppressant and moisture retaining properties of the mulch. However, additional labour
is expended when planting through a mulch containing a cardboard/newspaper layer, as
holes must be cut for each plant. Sowing seed through mulches containing a cardboard/
newspaper layer is impractical. Application of newspaper mulch in windy weather can be
facilitated by briefly pre-soaking the newspaper in water to increase its weight.

Carpet
• Synthetic carpet that is composed of artificial fibers may be removed after planting to pre-
vent fibers taking a long time to decompose, whereas carpet made from natural fibers may
be kept in place, blocking competition from weeds. Rain is absorbed by carpet and then
slowly released into the soil, reducing watering needs.

Colored Mulch
Some organic mulches are colored red, brown, black, and other colors. Isopropanolamine,
specifically 1-Amino-2-propanol or DOW™ monoisopropanolamine, may be used as a pigment
dispersant and color fastener in these mulches. Types of mulch which can be dyed include:
wood chips, bark chips (barkdust) and pine straw. Colored mulch is made by dyeing the mulch
in a water-based solution of colorant and chemical binder. When colored mulch first entered
the market, most formulas were suspected to contain toxic, heavy metals and other contami-
nates. Today, “current investigations indicate that mulch colorants pose no threat to people,
pets or the environment. The dyes currently used by the mulch and soil industry are similar
to those used in the cosmetic and other manufacturing industries (i.e., iron oxide),” as stated
by the Mulch and Soil Council. Colored mulch can be applied anywhere non-colored mulch is
used (such as large bedded areas or around plants) and features many of the same gardening
benefits as traditional mulch, such as improving soil productivity and retaining moisture. As
mulch decomposes, just as with non-colored mulch, more mulch may need to be added to
continue providing benefits to the soil and plants. However, if mulch is faded, spraying dye to
previously spread mulch in order to restore color is an option.

Anaerobic (Sour) Mulch


Mulch normally smells like freshly cut wood, but sometimes develops a toxicity that causes it to
smell like vinegar, ammonia, sulfur or silage. This happens when material with ample nitrogen
content is not rotated often enough and it forms pockets of increased decomposition. When this
occurs, the process may become anaerobic and produce these phytotoxic materials in small quan-
tities. Once exposed to the air, the process quickly reverts to an aerobic process, but these toxic
materials may be present for a period of time. If the mulch is placed around plants before the toxic-
ity has had a chance to dissipate, then the plants could very likely be damaged or killed depending

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226 Fundamentals of Weed Science

on their hardiness. Plants that are predominantly low to the ground or freshly planted are the most
susceptible, and the phytotoxicity may prevent germination of some seeds.

If sour mulch is applied and there is plant kill, the best thing to do is to water the mulch heavily.
Water dissipates the chemicals faster and refreshes the plants. Removing the offending mulch may
have little effect, because by the time plant kill is noticed, most of the toxicity is already dissipated.
While testing after plant kill will not likely turn up anything, a simple pH check may reveal high
acidity, in the range of 3.8 to 5.6 instead of the normal range of 6.0 to 7.2. Finally, placing a bit of
the offending mulch around another plant to check for plant kill will verify if the toxicity has de-
parted. If the new plant is also killed, then sour mulch is probably not the problem.

Groundcovers (Living Mulches)


Groundcovers are plants which grow close to the ground, under the main crop, to slow the devel-
opment of weeds and provide other benefits of mulch. They are usually fast-growing plants that
continue growing with the main crops. By contrast, cover crops are incorporated into the soil or
killed with herbicides. However, live mulches also may need to be mechanically or chemically
killed eventually to prevent competition with the main crop.

Some groundcovers can perform additional roles in the garden such as nitrogen fixation in the
case of clovers, dynamic accumulation of nutrients from the subsoil in the case of creeping comfrey
(Symphytum ibericum), and even food production in the case of Rubus tricolor.

On-site Mulch Production


Owing to the great bulk of mulch which is often required on a site, it is often impractical and ex-
pensive to source and import sufficient mulch materials. An alternative to importing mulch ma-
terials is to grow them on site in a “mulch garden” - an area of the site dedicated entirely to the
production of mulch which is then transferred to the growing area. Mulch gardens should be sited
as close as possible to the growing area so as to facilitate transfer of mulch materials.

Mulching (Composting) Over Unwanted Plants


Sufficient mulch over plants will destroy them, and may be more advantageous than using her-
bicide, cutting, mowing, pulling, raking, or tilling. The higher the temperature that this “mulch”
is composted, the quicker the reduction of undesirable materials. “Undesirable materials” may
include living seed, plant “trash”, as well as pathogens such as from animal feces, urine (e.g. han-
tavirus), fleas, lice, ticks, etc.

In some ways this improves the soil by attracting and feeding earthworms, and adding humus.
Earthworms “till” the soil, and their feces are among the best fertilizers and soil conditioners.

Urine may be toxic to plants if applied to growing areas undiluted.

Polypropylene and Polyethylene Mulch


Polypropylene mulch is made up of polypropylene polymers where polyethylene mulch is made

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Weed Control: Methods and Techniques 227

up of polyethylene polymers. These mulches are commonly used in many plastics. Polyethylene
is used mainly for weed reduction, where polypropylene is used mainly on perennials. This mulch
is placed on top of the soil and can be done by machine or hand with pegs to keep the mulch tight
against the soil. This mulch can prevent soil erosion, reduce weeding, conserve soil moisture, and
increase temperature of the soil. Ultimately this can reduce the amount of work a farmer may
have to do, and the amount of herbicides applied during the growing period. The black and clear
mulches capture sunlight and warm the soil increasing the growth rate. White and other reflective
colours will also warm the soil, but they do not suppress weeds as well. This mulch may require
other sources of obtaining water such as drip irrigation since it can reduce the amount of water
that reaches the soil. This mulch needs to be manually removed at the end of the season since when
it starts to break down it breaks down into smaller pieces. If the mulch is not removed before it
starts to break down eventually it will break down into ketones and aldehydes polluting the soil.
This mulch is technically biodegradable but does not break down into the same materials the more
natural biodegradable mulch does.

Biodegradable Mulch
Quality biodegradable mulches are made out of plant starches and sugars or polyester fibres.
These starches can come from plants such as wheat and corn. These mulch films may be a bit
more permeable allowing more water into the soil. This mulch can prevent soil erosion, reduce
weeding, conserve soil moisture, and increase temperature of the soil. Ultimately this can reduce
the amount of herbicides used and manual labour farmers may have to do throughout the growing
season. At the end of the season these mulches will start to break down from heat. Microorganisms
in the soil break down the mulch into two components, water and CO2, leaving no toxic residues
behind. This source of mulch is even less manual labour since it does not need to be removed at the
end of the season and can actually be tilled into the soil. With this mulch its important to take into
consideration that its mulch is more delicate then other kinds. It should be placed on a day which
is not too hot and with less tension then other synthetic mulches. These also can be placed by ma-
chine or hand but its ideal to have a more starchy mulch that will allow it to stick to the soil better.

Hoe (Tool)

A farmer using a hoe to keep weeds down in a vegetable garden.

A hoe is an ancient and versatile agricultural hand tool used to shape the soil, control weeds, clear

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228 Fundamentals of Weed Science

soil, and harvest root crops. Shaping the soil can be piling soil around the base of plants (hilling),
creating narrow furrows (drills) and shallow trenches for planting seeds and bulbs. Weed control
with a hoe can be by agitating the surface of the soil or by cutting foliage from the roots, and clear-
ing soil of old roots and crop residues. Hoes for digging and moving soil are used harvesting root
crops such as potatoes.

Types
There are many types of hoes of quite different appearances and purposes. Some can perform mul-
tiple functions. Others are intended for a specific use.

Cultivating tool pull or draw hoe

There are two main classes of agricultural hoe: draw hoes for shaping, and scuffle hoes for surface
weed control.

A draw hoe has the blade set at approximately a right angle to the handle. The user chops into the
ground and then pulls (draws) the blade towards them. Altering the angle of the handle can cause
the hoe to dig deeper or more shallowly as the hoe is pulled. A draw hoe can easily be used to cul-
tivate soil to a depth of several inches. A typical design of draw hoe, the “eye hoe”, has a ring in the
head through which the handle is fitted. This design has been used since Roman times.

A scuffle hoe is used to scrape the surface of the soil, and to loosen the top inch or so, and to cut
the roots, remove, and disrupt the growth of weeds efficiently. These are mainly of two different
designs: the Dutch hoe and the hoop hoe.

Eye hoe heads, some with sow-tooth (German: Sauzahn), Centro Etnográfico de Soutelo de
Montes, Pontevedra, Spain

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Weed Control: Methods and Techniques 229

Cultivating tool push or thrust hoe

The term ‘hand hoe’ most commonly refers to any type of light-weight, short-handled hoe, al-
though it may be used simply to contrast hand-held tools against animal or machine pulled tools.

Draw Hoes
• The typical farming and gardening hoe with a heavy, broad blade and a straight edge is
known as the grub hoe, grab hoe, pattern hoe, Italian hoe, azada or dago hoe (“dago” is an
ethnic slur referring to Italians, Spaniards, or Portuguese).

• Ridging hoes also known as warren or drill hoes are triangular (point-down) or heart-
shaped draw hoes particularly for creating narrow furrows (drills) and shallow trenches
for planting seeds and bulbs.

Hoedad (tree-planting tool) Kaibab National Forest, Arizona, USA

• Hoedads (also, “hoedags”) are hoe-like tools used for planting trees. According to Hartzell
(1987, p. 29), “The hoedag [was] originally called skindvic hoe... Hans Rasmussen, legend-
ary contractor and timber farm owner, is credited with having invented the curved, convex,
round-nosed hoedag blade which is widely used today” (emphasis added).

• The mortar hoe, a tool specific to hand mixing mortar and concrete, has the appearance of
a typical square-bladed draw hoe with the addition of large holes in the blade.

Scuffle Hoes
• The Dutch hoe is a design that is pushed or pulled through the soil to cut weeds just under
the surface. A Dutch hoe has a blade “sharp on every side so as to cut either forwards and
backwards”. The blade must be set in a plane slightly upwardly inclined to the dual axis of
the rod used as a handle stick. The user uses the handle to push the blade forward, forcing

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230 Fundamentals of Weed Science

it below the surface of the ground and maintaining it at a shallow depth in the surface layer
of soil by altering the angle of the handle whilst pushing. A push hoe can easily cultivate
and remove weeds etc. from the surface layer of the soil.

• The hoop hoe (also known as action, oscillating, hula, stirrup, pendulum weeder, or swivel
hoes) have a double-edge blade that bends around to form a rectangle attached to the han-
dle. Weeds are cut just below the soil surface as the blade is pushed & pulled through the
area. The back and forth motion is highly effective with cutting weeds in loose or breakable
soil. Widths of the blade typically range between three and seven inches. Its tool-head is a
loop of flat, sharpened strap metal. It is not as efficient as a draw hoe for moving soil.

• The collinear hoe has a narrow, razor-sharp blade which is used to slice weeds by skimming
it just under the surface of the soil with a sweeping motion; it is unsuitable for tasks like soil
moving and chopping. It was designed by Eliot Coleman in the late 1980s.

• The Swoe hoe is a modern one-sided cutting hoe - a variant of the Dutch hoe.

Other Hoes
Hoes resembling neither draw nor scuffle hoes include:

• Wheel hoes are, as the name suggests, a hoe or pair of hoes attached to one or more wheels.
The hoes are frequently interchangeable with other tools.

• Horse hoes, resembling small ploughs, were a favourite implement of agricultural pioneer
Jethro Tull, claiming in his book “Horse Hoeing Husbandry” that “the horse-hoe will, in
wide intervals, give wheat throughout all the stages of its life, as much nourishment as the
discreet hoer pleases”. The modern view is that, rather than nutrients being released, the
crop simply benefits from the removal of competing plants. The introduction of the horse
hoe, together with the better-known seed drill, brought about the great increase farming
productivity seen during the British Agricultural Revolution.

Fork-hoe depiction in Der Rebmann (the vine-dresser). Jost Amman, Das Ständebuch, 1568

• Fork hoes, (also known as prong hoes, tined hoes or bent forks) are hoes that have two
or more tines at right angles to the shaft. Their use is typically to loosen the soil, prior to
planting or sowing.

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Weed Control: Methods and Techniques 231

• Clam hoes, made for clam digging

• Adze hoes, with the basic hoe shape but heavier and stronger and with traditional uses in
trail making,

• Pacul or cangkul (hoes similar to adze hoe from Malaysia and Indonesia)

• Gang hoes for powered use (in use at least from 1887 to 1964).

History
Hoes are an ancient technology, predating the plough and perhaps preceded only by the digging
stick. In Sumerian mythology, the invention of the hoe was credited to Enlil, the chief of the coun-
cil of gods. The hand-plough (mr) was depicted in predynastic Egyptian art, and hoes are also
mentioned in ancient documents like the Code of Hammurabi (ca. 18th century BC) and the Book
of Isaiah (c. 8th century BC).

The human damage caused by long-term use of short-handled hoes, which required the user to
bend over from the waist to reach the ground, and caused permanent, crippling lower back pain to
farm workers, resulted, after struggle led by César Chávez with political help from Governor Jerry
Brown in the California Supreme Court declaring the short-handled hoe to be an unsafe hand tool
that was banned under California law in 1975.

‘Mr’ hand-plough, Protodynastic Period of Egypt (from the Scorpion Macehead)

• Ancient Greek iron hoe (Kerameikos Archaeological Museum)

A 2000-year-old iron Roman hoe blade

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232 Fundamentals of Weed Science

A Dutch hoe or push hoe; usually attached to a long hilt and handle

Archaeological Use
Over the past fifteen or twenty years, hoes have become increasingly popular tools for professional
archaeologists. While not as accurate as the traditional trowel, the hoe is an ideal tool for cleaning
relatively large open areas of archaeological interest. It is faster to use than a trowel, and produc-
es a much cleaner surface than an excavator bucket or shovel-scrape, and consequently on many
open-area excavations the once-common line of kneeling archaeologists trowelling backwards has
been replaced with a line of stooping archaeologists with hoes.

Herbicide
Herbicide(s), also commonly known as weedkillers, are chemical substances used to control un-
wanted plants. Selective herbicides control specific weed species, while leaving the desired crop
relatively unharmed, while non-selective herbicides (sometimes called “total weedkillers” in com-
mercial products) can be used to clear waste ground, industrial and construction sites, railways
and railway embankments as they kill all plant material with which they come into contact. Apart
from selective/non-selective, other important distinctions include persistence (also known as re-
sidual action: how long the product stays in place and remains active), means of uptake (whether
it is absorbed by above-ground foliage only, through the roots, or by other means), and mechanism
of action (how it works). Historically, products such as common salt and other metal salts were
used as herbicides, however these have gradually fallen out of favor and in some countries a num-
ber of these are banned due to their persistence in soil, and toxicity and groundwater contamina-
tion concerns. Herbicides have also been used in warfare and conflict.

Weeds controlled with herbicide

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Weed Control: Methods and Techniques 233

Modern herbicides are often synthetic mimics of natural plant hormones which interfere with
growth of the target plants. The term organic herbicide has come to mean herbicides intended for
organic farming; these are often less efficient and more costly than synthetic herbicides and are
based on natural materials. Some plants also produce their own natural herbicides, such as the
genus Juglans (walnuts), or the tree of heaven; such action of natural herbicides, and other related
chemical interactions, is called allelopathy. Due to herbicide resistance - a major concern in agri-
culture - a number of products also combine herbicides with different means of action.

In the US in 2007, about 83% of all herbicide usage, determined by weight applied, was in agri-
culture. In 2007, world pesticide expenditures totaled about $39.4 billion; herbicides were about
40% of those sales and constituted the biggest portion, followed by insecticides, fungicides, and
other types. Smaller quantities are used in forestry, pasture systems, and management of areas set
aside as wildlife habitat.

History
Prior to the widespread use of chemical herbicides, cultural controls, such as altering soil pH, sa-
linity, or fertility levels, were used to control weeds. Mechanical control (including tillage) was also
(and still is) used to control weeds.

First Herbicides

2,4-D, the first chemical herbicide, was discovered during the Second World War.

Although research into chemical herbicides began in the early 20th century, the first major break-
through was the result of research conducted in both the UK and the US during the Second World
War into the potential use of agents as biological weapons. The first modern herbicide, 2,4-D, was
first discovered and synthesized by W. G. Templeman at Imperial Chemical Industries. In 1940, he
showed that “Growth substances applied appropriately would kill certain broad-leaved weeds in
cereals without harming the crops.” By 1941, his team succeeded in synthesizing the chemical. In
the same year, Pokorny in the US achieved this as well.

Independently, a team under Juda Hirsch Quastel, working at the Rothamsted Experimental Station
made the same discovery. Quastel was tasked by the Agricultural Research Council (ARC) to discover
methods for improving crop yield. By analyzing soil as a dynamic system, rather than an inert sub-
stance, he was able to apply techniques such as perfusion. Quastel was able to quantify the influence
of various plant hormones, inhibitors and other chemicals on the activity of microorganisms in the
soil and assess their direct impact on plant growth. While the full work of the unit remained secret,
certain discoveries were developed for commercial use after the war, including the 2,4-D compound.

When it was commercially released in 1946, it triggered a worldwide revolution in agricultural out-

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234 Fundamentals of Weed Science

put and became the first successful selective herbicide. It allowed for greatly enhanced weed con-
trol in wheat, maize (corn), rice, and similar cereal grass crops, because it kills dicots (broadleaf
plants), but not most monocots (grasses). The low cost of 2,4-D has led to continued usage today,
and it remains one of the most commonly used herbicides in the world. Like other acid herbicides,
current formulations use either an amine salt (often trimethylamine) or one of many esters of the
parent compound. These are easier to handle than the acid.

Further Discoveries
The triazine family of herbicides, which includes atrazine, were introduced in the 1950s; they have
the current distinction of being the herbicide family of greatest concern regarding groundwater
contamination. Atrazine does not break down readily (within a few weeks) after being applied
to soils of above neutral pH. Under alkaline soil conditions, atrazine may be carried into the soil
profile as far as the water table by soil water following rainfall causing the aforementioned contam-
ination. Atrazine is thus said to have “carryover”, a generally undesirable property for herbicides.

Glyphosate (Roundup) was introduced in 1974 for nonselective weed control. Following the de-
velopment of glyphosate-resistant crop plants, it is now used very extensively for selective weed
control in growing crops. The pairing of the herbicide with the resistant seed contributed to the
consolidation of the seed and chemistry industry in the late 1990s.

Many modern chemical herbicides used in agriculture and gardening are specifically formulated to
decompose within a short period after application. This is desirable, as it allows crops and plants
to be planted afterwards, which could otherwise be affected by the herbicide. However, herbicides
with low residual activity (i.e., that decompose quickly) often do not provide season-long weed
control and do not ensure that weed roots are killed beneath construction and paving (and cannot
emerge destructively in years to come), therefore there remains a role for weedkiller with high
levels of persistence in the soil.

Terminology
Herbicides are classified/grouped in various ways e.g. according to the activity, timing of applica-
tion, method of application, mechanism of action, chemical family. This gives rise to a consider-
able level of terminology related to herbicides and their use.

Intended Outcome
• Control is the destruction of unwanted weeds, or the damage of them to the point where
they are no longer competitive with the crop.

• Suppression is incomplete control still providing some economic benefit, such as reduced
competition with the crop.

• Crop safety, for selective herbicides, is the relative absence of damage or stress to the crop.
Most selective herbicides cause some visible stress to crop plants.

• Defoliant, similar to herbicides, but designed to remove foliage (leaves) rather than kill the
plant.

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Weed Control: Methods and Techniques 235

Selectivity (All Plants or Specific Plants)


• Selective herbicides: They control or suppress certain plants without affecting the growth
of other plants species. Selectivity may be due to translocation, differential absorption,
physical (morphological) or physiological differences between plant species. 2,4-D, meco-
prop, dicamba control many broadleaf weeds but remain ineffective against turfgrasses.

• Non-selective herbicides: These herbicides are not specific in acting against certain plant
species and control all plant material with which they come into contact. They are used to
clear industrial sites, waste ground, railways and railway embankments. Paraquat, glufos-
inate, glyphosate are non-selective herbicides.

Timing of Application
• Preplant: Preplant herbicides are nonselective herbicides applied to soil before planting.
Some preplant herbicides may be mechanically incorporated into the soil. The objective for
incorporation is to prevent dissipation through photodecomposition and/or volatility. The
herbicides kill weeds as they grow through the herbicide treated zone. Volatile herbicides
have to be incorporated into the soil before planting the pasture. Agricultural crops grown
in soil treated with a preplant herbicide include tomatoes, corn, soybeans and strawberries.
Soil fumigants like metam-sodium and dazomet are in use as preplant herbicides.

• Preemergence: Preemergence herbicides are applied before the weed seedlings emerge
through the soil surface. Herbicides do not prevent weeds from germinating but they kill
weeds as they grow through the herbicide treated zone by affecting the cell division in the
emerging seedling. Dithopyr and pendimethalin are preemergence herbicides. Weeds that
have already emerged before application or activation are not affected by pre-herbicides as
their primary growing point escapes the treatment.

• Postemergence: These herbicides are applied after weed seedlings have emerged through
the soil surface. They can be foliar or root absorbed, selective or nonselective, contact or
systemic. Application of these herbicides is avoided during rain because the problem of be-
ing washed off to the soil makes it ineffective. 2,4-D is a selective, systemic, foliar absorbed
postemergence herbicide.

Method of Application
• Soil applied: Herbicides applied to the soil are usually taken up by the root or shoot of the
emerging seedlings and are used as preplant or preemergence treatment. Several factors
influence the effectiveness of soil-applied herbicides. Weeds absorb herbicides by both pas-
sive and active mechanism. Herbicide adsorption to soil colloids or organic matter often
reduces its amount available for weed absorption. Positioning of herbicide in correct layer
of soil is very important, which can be achieved mechanically and by rainfall. Herbicides on
the soil surface are subjected to several processes that reduce their availability. Volatility
and photolysis are two common processes that reduce the availability of herbicides. Many
soil applied herbicides are absorbed through plant shoots while they are still underground
leading to their death or injury. EPTC and trifluralin are soil applied herbicides.

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236 Fundamentals of Weed Science

• Foliar applied: These are applied to portion of the plant above the ground and are ab-
sorbed by exposed tissues. These are generally postemergence herbicides and can either be
translocated (systemic) throughout the plant or remain at specific site (contact). External
barriers of plants like cuticle, waxes, cell wall etc. affect herbicide absorption and action.
Glyphosate, 2,4-D and dicamba are foliar applied herbicide.

Persistence
• Residual activity: A herbicide is described as having low residual activity if it is neutralized
within a short time of application (within a few weeks or months) - typically this is due to
rainfall, or by reactions in the soil. A herbicide described as having high residual activity
will remains potent for a long term in the soil. For some compounds, the residual activity
can leave the ground almost permanently barren.

Mechanism of Action
Herbicides are often classified according to their site of action, because as a general rule, herbicides
within the same site of action class will produce similar symptoms on susceptible plants. Classifica-
tion based on site of action of herbicide is comparatively better as herbicide resistance management
can be handled more properly and effectively. Classification by mechanism of action (MOA) indi-
cates the first enzyme, protein, or biochemical step affected in the plant following application.

List of Mechanisms Found in Modern Herbicides


• ACCase inhibitors compounds kill grasses. Acetyl coenzyme A carboxylase (ACCase) is part
of the first step of lipid synthesis. Thus, ACCase inhibitors affect cell membrane production
in the meristems of the grass plant. The ACCases of grasses are sensitive to these herbi-
cides, whereas the ACCases of dicot plants are not.

• ALS inhibitors: the acetolactate synthase (ALS) enzyme (also known as acetohydroxyacid
synthase, or AHAS) is the first step in the synthesis of the branched-chain amino acids (va-
line, leucine, and isoleucine). These herbicides slowly starve affected plants of these amino
acids, which eventually leads to inhibition of DNA synthesis. They affect grasses and dicots
alike. The ALS inhibitor family includes various sulfonylureas (such as Flazasulfuron and
Metsulfuron-methyl), imidazolinones, triazolopyrimidines, pyrimidinyl oxybenzoates, and
sulfonylamino carbonyl triazolinones. The ALS biological pathway exists only in plants and
not animals, thus making the ALS-inhibitors among the safest herbicides.

• EPSPS inhibitors: The enolpyruvylshikimate 3-phosphate synthase enzyme EPSPS is used


in the synthesis of the amino acids tryptophan, phenylalanine and tyrosine. They affect
grasses and dicots alike. Glyphosate (Roundup) is a systemic EPSPS inhibitor inactivated
by soil contact.

• Synthetic auxins inaugurated the era of organic herbicides. They were discovered in the
1940s after a long study of the plant growth regulator auxin. Synthetic auxins mimic this
plant hormone. They have several points of action on the cell membrane, and are effective
in the control of dicot plants. 2,4-D is a synthetic auxin herbicide.

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Weed Control: Methods and Techniques 237

• Photosystem II inhibitors reduce electron flow from water to NADPH2+ at the pho-
tochemical step in photosynthesis. They bind to the Qb site on the D1 protein, and
prevent quinone from binding to this site. Therefore, this group of compounds causes
electrons to accumulate on chlorophyll molecules. As a consequence, oxidation reac-
tions in excess of those normally tolerated by the cell occur, and the plant dies. The
triazine herbicides (including atrazine) and urea derivatives (diuron) are photosystem
II inhibitors.

• Photosystem I inhibitors steal electrons from the normal pathway through FeS to Fdx to
NADP leading to direct discharge of electrons on oxygen. As a result, reactive oxygen spe-
cies are produced and oxidation reactions in excess of those normally tolerated by the cell
occur, leading to plant death. Bipyridinium herbicides (such as diquat and paraquat) in-
hibit the Fe-S – Fdx step of that chain, while diphenyl ether herbicides (such as nitrofen,
nitrofluorfen, and acifluorfen) inhibit the Fdx – NADP step.

• HPPD inhibitors inhibit 4-Hydroxyphenylpyruvate dioxygenase, which are involved in ty-


rosine breakdown. Tyrosine breakdown products are used by plants to make carotenoids,
which protect chlorophyll in plants from being destroyed by sunlight. If this happens, the
plants turn white due to complete loss of chlorophyll, and the plants die. Mesotrione and
sulcotrione are herbicides in this class; a drug, nitisinone, was discovered in the course of
developing this class of herbicides.

Herbicide Group (Labeling)


One of the most important methods for preventing, delaying, or managing resistance is to
reduce the reliance on a single herbicide mode of action. To do this, farmers must know the
mode of action for the herbicides they intend to use, but the relatively complex nature of plant
biochemistry makes this difficult to determine. Attempts were made to simplify the under-
standing of herbicide mode of action by developing a classification system that grouped herbi-
cides by mode of action. Eventually the Herbicide Resistance Action Committee (HRAC) and
the Weed Science Society of America (WSSA) developed a classification system. The WSSA and
HRAC systems differ in the group designation. Groups in the WSSA and the HRAC systems are
designated by numbers and letters, respectively. The goal for adding the “Group” classification
and mode of action to the herbicide product label is to provide a simple and practical approach
to deliver the information to users. This information will make it easier to develop educational
material that is consistent and effective. It should increase user’s awareness of herbicide mode
of action and provide more accurate recommendations for resistance management. Another
goal is to make it easier for users to keep records on which herbicide mode of actions are being
used on a particular field from year to year.

Chemical Family
Detailed investigations on chemical structure of the active ingredients of the registered herbicides
showed that some moieties (moiety is a part of a molecule that may include either whole func-
tional groups or parts of functional groups as substructures; a functional group has similar chem-
ical properties whenever it occurs in different compounds) have the same mechanisms of action.

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238 Fundamentals of Weed Science

According to Forouzesh et al. 2015, these moieties have been assigned to the names of chemical
families and active ingredients are then classified within the chemical families accordingly. Know-
ing about herbicide chemical family grouping could serve as a short-term strategy for managing
resistance to site of action.

Use and Application

Herbicides being sprayed from the spray arms of a tractor in North Dakota.

Most herbicides are applied as water-based sprays using ground equipment. Ground equipment
varies in design, but large areas can be sprayed using self-propelled sprayers equipped with long
booms, of 60 to 120 feet (18 to 37 m) with spray nozzles spaced every 20–30 inches (510–760 mm)
apart. Towed, handheld, and even horse-drawn sprayers are also used. On large areas, herbicides
may also at times be applied aerially using helicopters or airplanes, or through irrigation systems
(known as chemigation).

A further method of herbicide application developed around 2010, involves ridding the soil of its
active weed seed bank rather than just killing the weed. This can successfully treat annual plants
but not perennials. Researchers at the Agricultural Research Service found that the application of
herbicides to fields late in the weeds’ growing season greatly reduces their seed production, and
therefore fewer weeds will return the following season. Because most weeds are annuals, their
seeds will only survive in soil for a year or two, so this method will be able to destroy such weeds
after a few years of herbicide application.

Weed-wiping may also be used, where a wick wetted with herbicide is suspended from a boom
and dragged or rolled across the tops of the taller weed plants. This allows treatment of taller
grassland weeds by direct contact without affecting related but desirable shorter plants in
the grassland sward beneath. The method has the benefit of avoiding spray drift. In Wales, a
scheme offering free weed-wiper hire was launched in 2015 in an effort to reduce the levels of
MCPA in water courses.

Misuse and Misapplication


Herbicide volatilisation or spray drift may result in herbicide affecting neighboring fields or
plants, particularly in windy conditions. Sometimes, the wrong field or plants may be sprayed
due to error.

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Weed Control: Methods and Techniques 239

Use Politically, Militarily, and in Conflict


Health and Environmental Effects
Herbicides have widely variable toxicity in addition to acute toxicity from occupational exposure
levels.

Some herbicides cause a range of health effects ranging from skin rashes to death. The pathway
of attack can arise from intentional or unintentional direct consumption, improper application
resulting in the herbicide coming into direct contact with people or wildlife, inhalation of aerial
sprays, or food consumption prior to the labeled preharvest interval. Under some conditions, cer-
tain herbicides can be transported via leaching or surface runoff to contaminate groundwater or
distant surface water sources. Generally, the conditions that promote herbicide transport include
intense storm events (particularly shortly after application) and soils with limited capacity to ad-
sorb or retain the herbicides. Herbicide properties that increase likelihood of transport include
persistence (resistance to degradation) and high water solubility.

Phenoxy herbicides are often contaminated with dioxins such as TCDD; research has suggested
such contamination results in a small rise in cancer risk after occupational exposure to these her-
bicides. Triazine exposure has been implicated in a likely relationship to increased risk of breast
cancer, although a causal relationship remains unclear.

Herbicide manufacturers have at times made false or misleading claims about the safety of their
products. Chemical manufacturer Monsanto Company agreed to change its advertising after pres-
sure from New York attorney general Dennis Vacco; Vacco complained about misleading claims
that its spray-on glyphosate-based herbicides, including Roundup, were safer than table salt and
“practically non-toxic” to mammals, birds, and fish (though proof that this was ever said is hard to
find). Roundup is toxic and has resulted in death after being ingested in quantities ranging from
85 to 200 ml, although it has also been ingested in quantities as large as 500 ml with only mild
or moderate symptoms. The manufacturer of Tordon 101 (Dow AgroSciences, owned by the Dow
Chemical Company) has claimed Tordon 101 has no effects on animals and insects, in spite of evi-
dence of strong carcinogenic activity of the active ingredient Picloram in studies on rats.

The risk of Parkinson’s disease has been shown to increase with occupational exposure to herbi-
cides and pesticides. The herbicide paraquat is suspected to be one such factor.

All commercially sold, organic and nonorganic herbicides must be extensively tested prior to ap-
proval for sale and labeling by the Environmental Protection Agency. However, because of the
large number of herbicides in use, concern regarding health effects is significant. In addition to
health effects caused by herbicides themselves, commercial herbicide mixtures often contain other
chemicals, including inactive ingredients, which have negative impacts on human health.

Ecological Effects
Commercial herbicide use generally has negative impacts on bird populations, although the im-
pacts are highly variable and often require field studies to predict accurately. Laboratory studies
have at times overestimated negative impacts on birds due to toxicity, predicting serious problems
that were not observed in the field. Most observed effects are due not to toxicity, but to habitat

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240 Fundamentals of Weed Science

changes and the decreases in abundance of species on which birds rely for food or shelter. Herbi-
cide use in silviculture, used to favor certain types of growth following clearcutting, can cause sig-
nificant drops in bird populations. Even when herbicides which have low toxicity to birds are used,
they decrease the abundance of many types of vegetation on which the birds rely. Herbicide use
in agriculture in Britain has been linked to a decline in seed-eating bird species which rely on the
weeds killed by the herbicides. Heavy use of herbicides in neotropical agricultural areas has been
one of many factors implicated in limiting the usefulness of such agricultural land for wintering
migratory birds.

Frog populations may be affected negatively by the use of herbicides as well. While some studies
have shown that atrazine may be a teratogen, causing demasculinization in male frogs, the U.S.
Environmental Protection Agency (EPA) and its independent Scientific Advisory Panel (SAP) ex-
amined all available studies on this topic and concluded that “atrazine does not adversely affect
amphibian gonadal development based on a review of laboratory and field studies.”

Scientific Uncertainty of Full Extent of Herbicide Effects


The health and environmental effects of many herbicides is unknown, and even the scientific com-
munity often disagrees on the risk. For example, a 1995 panel of 13 scientists reviewing studies
on the carcinogenicity of 2,4-D had divided opinions on the likelihood 2,4-D causes cancer in hu-
mans. As of 1992 , studies on phenoxy herbicides were too few to accurately assess the risk of many
types of cancer from these herbicides, even though evidence was stronger that exposure to these
herbicides is associated with increased risk of soft tissue sarcoma and non-Hodgkin lymphoma.
Furthermore, there is some suggestion that herbicides can play a role in sex reversal of certain
organisms that experience temperature-dependent sex determination, which could theoretically
alter sex ratios.

Resistance
Weed resistance to herbicides has become a major concern in crop production worldwide. Re-
sistance to herbicides is often attributed to lack of rotational programmes of herbicides and to
continuous applications of herbicides with the same sites of action. Thus, a true understanding of
the sites of action of herbicides is essential for strategic planning of herbicide-based weed control.

Plants have developed resistance to atrazine and to ALS-inhibitors, and more recently, to glypho-
sate herbicides. Marestail is one weed that has developed glyphosate resistance. Glyphosate-resis-
tant weeds are present in the vast majority of soybean, cotton and corn farms in some U.S. states.
Weeds that can resist multiple other herbicides are spreading. Few new herbicides are near com-
mercialization, and none with a molecular mode of action for which there is no resistance. Because
most herbicides could not kill all weeds, farmers rotated crops and herbicides to stop resistant
weeds. During its initial years, glyphosate was not subject to resistance and allowed farmers to
reduce the use of rotation.

A family of weeds that includes waterhemp (Amaranthus rudis) is the largest concern. A 2008-9
survey of 144 populations of waterhemp in 41 Missouri counties revealed glyphosate resistance in
69%. Weeds from some 500 sites throughout Iowa in 2011 and 2012 revealed glyphosate resistance
in approximately 64% of waterhemp samples. The use of other killers to target “residual” weeds has

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Weed Control: Methods and Techniques 241

become common, and may be sufficient to have stopped the spread of resistance From 2005 through
2010 researchers discovered 13 different weed species that had developed resistance to glyphosate.
But since then only two more have been discovered. Weeds resistant to multiple herbicides with
completely different biological action modes are on the rise. In Missouri, 43% of samples were re-
sistant to two different herbicides; 6% resisted three; and 0.5% resisted four. In Iowa 89% of water-
hemp samples resist two or more herbicides, 25% resist three, and 10% resist five.

For southern cotton, herbicide costs has climbed from between $50 and $75 per hectare a few
years ago to about $370 per hectare in 2013. Resistance is contributing to a massive shift away
from growing cotton; over the past few years, the area planted with cotton has declined by 70% in
Arkansas and by 60% in Tennessee. For soybeans in Illinois, costs have risen from about $25 to
$160 per hectare.

Dow, Bayer CropScience, Syngenta and Monsanto are all developing seed varieties resistant to
herbicides other than glyphosate, which will make it easier for farmers to use alternative weed
killers. Even though weeds have already evolved some resistance to those herbicides, Powles says
the new seed-and-herbicide combos should work well if used with proper rotation.

Biochemistry of Resistance
Resistance to herbicides can be based on one of the following biochemical mechanisms:

• Target-site resistance: This is due to a reduced (or even lost) ability of the herbicide to bind
to its target protein. The effect usually relates to an enzyme with a crucial function in a met-
abolic pathway, or to a component of an electron-transport system. Target-site resistance
may also be caused by an overexpression of the target enzyme (via gene amplification or
changes in a gene promoter).

• Non-target-site resistance: This is caused by mechanisms that reduce the amount of herbi-
cidal active compound reaching the target site. One important mechanism is an enhanced
metabolic detoxification of the herbicide in the weed, which leads to insufficient amounts
of the active substance reaching the target site. A reduced uptake and translocation, or
sequestration of the herbicide, may also result in an insufficient herbicide transport to the
target site.

• Cross-resistance: In this case, a single resistance mechanism causes resistance to several


herbicides. The term target-site cross-resistance is used when the herbicides bind to the
same target site, whereas non-target-site cross-resistance is due to a single non-target-site
mechanism (e.g., enhanced metabolic detoxification) that entails resistance across herbi-
cides with different sites of action.

• Multiple resistance: In this situation, two or more resistance mechanisms are present with-
in individual plants, or within a plant population.

Resistance Management
Worldwide experience has been that farmers tend to do little to prevent herbicide resistance de-
veloping, and only take action when it is a problem on their own farm or neighbor’s. Careful obser-

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242 Fundamentals of Weed Science

vation is important so that any reduction in herbicide efficacy can be detected. This may indicate
evolving resistance. It is vital that resistance is detected at an early stage as if it becomes an acute,
whole-farm problem, options are more limited and greater expense is almost inevitable. Table 1
lists factors which enable the risk of resistance to be assessed. An essential pre-requisite for con-
firmation of resistance is a good diagnostic test. Ideally this should be rapid, accurate, cheap and
accessible. Many diagnostic tests have been developed, including glasshouse pot assays, petri dish
assays and chlorophyll fluorescence. A key component of such tests is that the response of the
suspect population to a herbicide can be compared with that of known susceptible and resistant
standards under controlled conditions. Most cases of herbicide resistance are a consequence of
the repeated use of herbicides, often in association with crop monoculture and reduced cultivation
practices. It is necessary, therefore, to modify these practices in order to prevent or delay the onset
of resistance or to control existing resistant populations. A key objective should be the reduction
in selection pressure. An integrated weed management (IWM) approach is required, in which as
many tactics as possible are used to combat weeds. In this way, less reliance is placed on herbicides
and so selection pressure should be reduced.

Optimising herbicide input to the economic threshold level should avoid the unnecessary use of
herbicides and reduce selection pressure. Herbicides should be used to their greatest potential by
ensuring that the timing, dose, application method, soil and climatic conditions are optimal for good
activity. In the UK, partially resistant grass weeds such as Alopecurus myosuroides (blackgrass) and
Avena spp. (wild oat) can often be controlled adequately when herbicides are applied at the 2-3 leaf
stage, whereas later applications at the 2-3 tiller stage can fail badly. Patch spraying, or applying
herbicide to only the badly infested areas of fields, is another means of reducing total herbicide use.

Table 1. Agronomic factors influencing the risk of herbicide resistance development

Factor Low risk High risk

Cropping system Good rotation Crop monoculture

Cultivation system Annual ploughing Continuous minimum tillage

Weed control Cultural only Herbicide only

Herbicide use Many modes of action Single modes of action

Control in previous years Excellent Poor

Weed infestation Low High

Resistance in vicinity Unknown Common

Approaches to Treating Resistant Weeds


Alternative Herbicides
When resistance is first suspected or confirmed, the efficacy of alternatives is likely to be the first
consideration. The use of alternative herbicides which remain effective on resistant populations
can be a successful strategy, at least in the short term. The effectiveness of alternative herbicides
will be highly dependent on the extent of cross-resistance. If there is resistance to a single group
of herbicides, then the use of herbicides from other groups may provide a simple and effective

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Weed Control: Methods and Techniques 243

solution, at least in the short term. For example, many triazine-resistant weeds have been readily
controlled by the use of alternative herbicides such as dicamba or glyphosate. If resistance extends
to more than one herbicide group, then choices are more limited. It should not be assumed that
resistance will automatically extend to all herbicides with the same mode of action, although it is
wise to assume this until proved otherwise. In many weeds the degree of cross-resistance between
the five groups of ALS inhibitors varies considerably. Much will depend on the resistance mech-
anisms present, and it should not be assumed that these will necessarily be the same in different
populations of the same species. These differences are due, at least in part, to the existence of dif-
ferent mutations conferring target site resistance. Consequently, selection for different mutations
may result in different patterns of cross-resistance. Enhanced metabolism can affect even close-
ly related herbicides to differing degrees. For example, populations of Alopecurus myosuroides
(blackgrass) with an enhanced metabolism mechanism show resistance to pendimethalin but not
to trifluralin, despite both being dinitroanilines. This is due to differences in the vulnerability of
these two herbicides to oxidative metabolism. Consequently, care is needed when trying to predict
the efficacy of alternative herbicides.

Mixtures and Sequences


The use of two or more herbicides which have differing modes of action can reduce the selection
for resistant genotypes. Ideally, each component in a mixture should:

• Be active at different target sites

• Have a high level of efficacy

• Be detoxified by different biochemical pathways

• Have similar persistence in the soil (if it is a residual herbicide)

• Exert negative cross-resistance

• Synergise the activity of the other component

No mixture is likely to have all these attributes, but the first two listed are the most important. There
is a risk that mixtures will select for resistance to both components in the longer term. One practical
advantage of sequences of two herbicides compared with mixtures is that a better appraisal of the
efficacy of each herbicide component is possible, provided that sufficient time elapses between each
application. A disadvantage with sequences is that two separate applications have to be made and it
is possible that the later application will be less effective on weeds surviving the first application. If
these are resistant, then the second herbicide in the sequence may increase selection for resistant in-
dividuals by killing the susceptible plants which were damaged but not killed by the first application,
but allowing the larger, less affected, resistant plants to survive. This has been cited as one reason
why ALS-resistant Stellaria media has evolved in Scotland recently (2000), despite the regular use
of a sequence incorporating mecoprop, a herbicide with a different mode of action.

Herbicide Rotations
Rotation of herbicides from different chemical groups in successive years should reduce selection

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244 Fundamentals of Weed Science

for resistance. This is a key element in most resistance prevention programmes. The value of this
approach depends on the extent of cross-resistance, and whether multiple resistance occurs owing
to the presence of several different resistance mechanisms. A practical problem can be the lack of
awareness by farmers of the different groups of herbicides that exist. In Australia a scheme has
been introduced in which identifying letters are included on the product label as a means of en-
abling farmers to distinguish products with different modes of action.

Farming Practices and Resistance: a Case Study


Herbicide resistance became a critical problem in Australian agriculture, after many Australian
sheep farmers began to exclusively grow wheat in their pastures in the 1970s. Introduced varieties
of ryegrass, while good for grazing sheep, compete intensely with wheat. Ryegrasses produce so
many seeds that, if left unchecked, they can completely choke a field. Herbicides provided excel-
lent control, while reducing soil disrupting because of less need to plough. Within little more than
a decade, ryegrass and other weeds began to develop resistance. In response Australian farmers
changed methods. By 1983, patches of ryegrass had become immune to Hoegrass, a family of her-
bicides that inhibit an enzyme called acetyl coenzyme A carboxylase.

Ryegrass populations were large, and had substantial genetic diversity, because farmers had plant-
ed many varieties. Ryegrass is cross-pollinated by wind, so genes shuffle frequently. To control
its distribution farmers sprayed inexpensive Hoegrass, creating selection pressure. In addition,
farmers sometimes diluted the herbicide in order to save money, which allowed some plants to
survive application. When resistance appeared farmers turned to a group of herbicides that block
acetolactate synthase. Once again, ryegrass in Australia evolved a kind of “cross-resistance” that
allowed it to rapidly break down a variety of herbicides. Four classes of herbicides become ineffec-
tive within a few years. In 2013 only two herbicide classes, called Photosystem II and long-chain
fatty acid inhibitors, were effective against ryegrass.

List of Common Herbicides


Synthetic Herbicides
• 2,4-D is a broadleaf herbicide in the phenoxy group used in turf and no-till field crop
production. Now, it is mainly used in a blend with other herbicides to allow lower rates
of herbicides to be used; it is the most widely used herbicide in the world, and third
most commonly used in the United States. It is an example of synthetic auxin (plant
hormone).

• Aminopyralid is a broadleaf herbicide in the pyridine group, used to control weeds on grass-
land, such as docks, thistles and nettles. It is notorious for its ability to persist in compost.

• Atrazine, a triazine herbicide, is used in corn and sorghum for control of broadleaf weeds
and grasses. Still used because of its low cost and because it works well on a broad spectrum
of weeds common in the US corn belt, atrazine is commonly used with other herbicides to
reduce the overall rate of atrazine and to lower the potential for groundwater contamina-
tion; it is a photosystem II inhibitor.

• Clopyralid is a broadleaf herbicide in the pyridine group, used mainly in turf, rangeland,

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Weed Control: Methods and Techniques 245

and for control of noxious thistles. Notorious for its ability to persist in compost, it is an-
other example of synthetic auxin.

• Dicamba, a postemergent broadleaf herbicide with some soil activity, is used on turf and
field corn. It is another example of a synthetic auxin.

• Glufosinate ammonium, a broad-spectrum contact herbicide, is used to control weeds after


the crop emerges or for total vegetation control on land not used for cultivation.

• Fluazifop (Fuselade Forte), a post emergence, foliar absorbed, translocated grass-selective


herbicide with little residual action. It is used on a very wide range of broad leaved crops
for control of annual and perennial grasses.

• Fluroxypyr, a systemic, selective herbicide, is used for the control of broad-leaved weeds
in small grain cereals, maize, pastures, rangeland and turf. It is a synthetic auxin. In cereal
growing, fluroxypyr’s key importance is control of cleavers, Galium aparine. Other key
broadleaf weeds are also controlled.

• Glyphosate, a systemic nonselective herbicide, is used in no-till burndown and for weed
control in crops genetically modified to resist its effects. It is an example of an EPSPs in-
hibitor.

• Imazapyr a nonselective herbicide, is used for the control of a broad range of weeds, in-
cluding terrestrial annual and perennial grasses and broadleaf herbs, woody species, and
riparian and emergent aquatic species.

• Imazapic, a selective herbicide for both the pre- and postemergent control of some annual
and perennial grasses and some broadleaf weeds, kills plants by inhibiting the production
of branched chain amino acids (valine, leucine, and isoleucine), which are necessary for
protein synthesis and cell growth.

• Imazamox, an imidazolinone manufactured by BASF for postemergence application that is an


acetolactate synthase (ALS) inhibitor. Sold under trade names Raptor, Beyond, and Clearcast.

• Linuron is a nonselective herbicide used in the control of grasses and broadleaf weeds. It
works by inhibiting photosynthesis.

• MCPA (2-methyl-4-chlorophenoxyacetic acid) is a phenoxy herbicide selective for broad-


leaf plants and widely used in cereals and pasture.

• Metolachlor is a pre-emergent herbicide widely used for control of annual grasses in corn
and sorghum; it has displaced some of the atrazine in these uses.

• Paraquat is a nonselective contact herbicide used for no-till burndown and in aerial de-
struction of marijuana and coca plantings. It is more acutely toxic to people than any other
herbicide in widespread commercial use.

• Pendimethalin, a pre-emergent herbicide, is widely used to control annual grasses and


some broad-leaf weeds in a wide range of crops, including corn, soybeans, wheat, cotton,
many tree and vine crops, and many turfgrass species.

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246 Fundamentals of Weed Science

• Picloram, a pyridine herbicide, mainly is used to control unwanted trees in pastures and
edges of fields. It is another synthetic auxin.

• Sodium chlorate (disused/banned in some countries), a nonselective herbicide, is consid-


ered phytotoxic to all green plant parts. It can also kill through root absorption.

• Triclopyr, a systemic, foliar herbicide in the pyridine group, is used to control broadleaf
weeds while leaving grasses and conifers unaffected.

• Several sulfonylureas, including Flazasulfuron and Metsulfuron-methyl, which act as ALS


inhibitors and in some cases are taken up from the soil via the roots.

Organic Herbicides
Recently, the term “organic” has come to imply products used in organic farming. Under this defi-
nition, an organic herbicide is one that can be used in a farming enterprise that has been classified
as organic. Commercially sold organic herbicides are expensive and may not be affordable for
commercial farming. Depending on the application, they may be less effective than synthetic her-
bicides and are generally used along with cultural and mechanical weed control practices.

Homemade organic herbicides include:

• Corn gluten meal (CGM) is a natural pre-emergence weed control used in turfgrass, which
reduces germination of many broadleaf and grass weeds.

• Vinegar is effective for 5–20% solutions of acetic acid, with higher concentrations most
effective, but it mainly destroys surface growth, so respraying to treat regrowth is needed.
Resistant plants generally succumb when weakened by respraying.

• Steam has been applied commercially, but is now considered uneconomical and inade-
quate. It controls surface growth but not underground growth and so respraying to treat
regrowth of perennials is needed.

• Flame is considered more effective than steam, but suffers from the same difficulties.

• D-limonene (citrus oil) is a natural degreasing agent that strips the waxy skin or cuticle
from weeds, causing dehydration and ultimately death.

• Saltwater or salt applied in appropriate strengths to the rootzone will kill most plants.

• Monocerin produced by certain fungi will kill certain weeds such as Johnson grass.

Of Historical Interest and Other


• 2,4,5-Trichlorophenoxyacetic acid (2,4,5-T) was a widely used broadleaf herbicide until
being phased out starting in the late 1970s. While 2,4,5-T itself is of only moderate toxicity,
the manufacturing process for 2,4,5-T contaminates this chemical with trace amounts of
2,3,7,8-tetrachlorodibenzo-p-dioxin (TCDD). TCDD is extremely toxic to humans. With
proper temperature control during production of 2,4,5-T, TCDD levels can be held to about
.005 ppm. Before the TCDD risk was well understood, early production facilities lacked

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Weed Control: Methods and Techniques 247

proper temperature controls. Individual batches tested later were found to have as much as
60 ppm of TCDD. 2,4,5-T was withdrawn from use in the USA in 1983, at a time of height-
ened public sensitivity about chemical hazards in the environment. Public concern about
dioxins was high, and production and use of other (non-herbicide) chemicals potentially
containing TCDD contamination was also withdrawn. These included pentachlorophenol
(a wood preservative) and PCBs (mainly used as stabilizing agents in transformer oil).
Some feel that the 2,4,5-T withdrawal was not based on sound science. 2,4,5-T has since
largely been replaced by dicamba and triclopyr.

• Agent Orange was a herbicide blend used by the British military during the Malayan Emer-
gency and the U.S. military during the Vietnam War between January 1965 and April 1970
as a defoliant. It was a 50/50 mixture of the n-butyl esters of 2,4,5-T and 2,4-D. Because
of TCDD contamination in the 2,4,5-T component, it has been blamed for serious illnesses
in many people who were exposed to it. However, research on populations exposed to its
dioxin contaminant have been inconsistent and inconclusive.

• Diesel, and other heavy oil derivatives, are known to be informally used at times, but are
usually banned for this purpose.

Glyphosate
Glyphosate (N-(phosphonomethyl)glycine) is a broad-spectrum systemic herbicide and crop des-
iccant. It is an organophosphorus compound, specifically a phosphonate. It is used to kill weeds,
especially annual broadleaf weeds and grasses that compete with crops. It was discovered to be
an herbicide by Monsanto chemist John E. Franz in 1970. Monsanto brought it to market in 1974
under the trade name Roundup, and Monsanto’s last commercially relevant United States patent
expired in 2000.

Farmers quickly adopted glyphosate, especially after Monsanto introduced glyphosate-resistant


Roundup Ready crops, enabling farmers to kill weeds without killing their crops. In 2007, gly-
phosate was the most used herbicide in the United States’ agricultural sector and the second-most
used in home and garden, government and industry, and commerce. By 2016 there was a 100-fold
increase from the late 1970s in the frequency of applications and volumes of glyphosate-based
herbicides (GBHs) applied, partly in response to the unprecedented global emergence and spread
of glyphosate-resistant weeds.

Glyphosate is absorbed through foliage, and minimally through roots, and transported to growing
points. It inhibits a plant enzyme involved in the synthesis of three aromatic amino acids: tyrosine,
tryptophan, and phenylalanine. Therefore, it is effective only on actively growing plants and is not
effective as a pre-emergence herbicide. An increasing number of crops have been genetically engi-
neered to be tolerant of glyphosate (e.g. Roundup Ready soybean, the first Roundup Ready crop,
also created by Monsanto) which allows farmers to use glyphosate as a postemergence herbicide
against weeds. The development of glyphosate resistance in weed species is emerging as a costly
problem. While glyphosate and formulations such as Roundup have been approved by regulatory
bodies worldwide, concerns about their effects on humans and the environment persist.

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248 Fundamentals of Weed Science

Many regulatory and scholarly reviews have evaluated the relative toxicity of glyphosate as an
herbicide. The German Federal Institute for Risk Assessment toxicology review in 2013 found
that “the available data is contradictory and far from being convincing” with regard to correlations
between exposure to glyphosate formulations and risk of various cancers, including non-Hodgkin
lymphoma (NHL). A meta-analysis published in 2014 identified an increased risk of NHL in work-
ers exposed to glyphosate formulations. In March 2015 the World Health Organization’s Interna-
tional Agency for Research on Cancer classified glyphosate as “probably carcinogenic in humans”
(category 2A) based on epidemiological studies, animal studies, and in vitro studies. In November,
2015, the European Food Safety Authority published an updated assessment report on glyphosate,
concluding that “the substance is unlikely to be genotoxic (i.e. damaging to DNA) or to pose a
carcinogenic threat to humans.” Furthermore, the final report clarified that while other, probably
carcinogenic, glyphosate-containing formulations may exist, studies “that look solely at the active
substance glyphosate do not show this effect.” In May 2016, the Joint FAO/WHO Meeting on Pes-
ticide Residues concluded that “glyphosate is unlikely to pose a carcinogenic risk to humans from
exposure through the diet”, even at doses as high as 2,000 mg/kg body weight orally.

Discovery
Glyphosate was first synthesized in 1950 by Swiss chemist Henry Martin, who worked for the Swiss
company Cilag. The work was never published. Stauffer Chemical patented the agent as a chem-
ical chelator in 1964 as it binds and removes minerals such as calcium, magnesium, manganese,
copper, and zinc.

Somewhat later, glyphosate was independently discovered at Monsanto in 1970. Monsanto chem-
ists had synthesized about 100 analogs of aminomethylphosphonic acid as potential water-soft-
ening agents. Two were found to have weak herbicidal activity, and John E. Franz, a chemist at
Monsanto, was asked to try to make analogs with stronger herbicidal activity. Glyphosate was the
third analog he made. Franz received the National Medal of Technology in 1987 and the Perkin
Medal for Applied Chemistry in 1990 for his discoveries.

In 2008, USDA-ARS scientist Stephen O. Duke and Stephen B. Powles, an Australian weed expert
— described glyphosate as a “virtually ideal” herbicide. In 2010 Powles stated: “glyphosate is a one
in a 100-year discovery that is as important for reliable global food production as penicillin is for
battling disease.”

Chemistry
Glyphosate is an aminophosphonic analogue of the natural amino acid glycine, and like all amino
acids, exists in different ionic states depending on pH. Both the phosphonic acid and carboxylic
acid moieties can be ionised and the amine group can be protonated and the substance exists as
a series of zwitterions. Glyphosate is soluble in water to 12 g/l at room temperature. The original
synthetic approach to glyphosate involved the reaction of phosphorus trichloride with formalde-
hyde followed by hydrolysis to yield a phosphonate. Glycine is then reacted with this phosphonate
to yield glyphosate, and its name is taken as a contraction of the compounds used in this synthesis
- viz. glycine and a phosphonate.

PCl3 + H2CO → Cl2P(=O)-CH2Cl

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Weed Control: Methods and Techniques 249

Cl2P(=O)-CH2Cl + 2 H2O → (HO)2P(=O)-CH2Cl + 2 HCl

(HO)2P(=O)-CH2Cl + H2N-CH2-COOH → (HO)2P(=O)-CH2-NH-CH2-COOH + HCl

The main deactivation path for glyphosate is hydrolysis to aminomethylphosphonic acid.

Industrial Synthesis
Two main approaches are used to synthesize glyphosate industrially. The first is to react imino-
diacetic acid with phosphorous acid and hydrochloric acid (sometimes formed in situ by addition
of phosphorus trichloride) via a modified Mannich reaction. Oxidation then leads to the desired
glyphosate product. Iminodiacetic acid is usually prepared on-site, such as by reaction of chloro-
acetic acid with ammonia and calcium hydroxide to produce the calcium iminodiacetate salt and
then acidification of the product.

The chloroacetic acid approach is less efficient than other iminodiacetic acid approaches, owing to
the production of calcium chloride waste and decreased yield. When hydrogen cyanide is readily
available as a by-product (say), an alternative approach is to use iminodiacetonitrile, HN(CH2CN)2,
and diethanolamine is also a suitable starting material.

The second involves the use of dimethyl phosphite in a one-pot synthesis. Glycine and paraformal-
dehyde are reacted in a suitable organic solvent (typically triethylamine and methanol) to produce
bishydroxymethylglycine, (HOCH2)2NCH2COOH. Dimethyl phosphite is then introduced and
work-up with hydrochloric acid cleaves the hydroxymethyl group from the nitrogen atom whilst
heating speeds the hydrolysis of both phosphate ester linkages.

This synthetic approach is responsible for a substantial portion of the production of glyphosate in
China, with considerable work having gone into recycling the triethylamine and methanol. Prog-
ress has also been made in attempting to eliminate the need for triethylamine altogether.

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250 Fundamentals of Weed Science

Mode of Action
Glyphosate kills plants by interfering with the synthesis of the aromatic amino acids phenylalanine,
tyrosine, and tryptophan. It does this by inhibiting the enzyme 5-enolpyruvylshikimate-3-phos-
phate synthase (EPSPS), which catalyzes the reaction of shikimate-3-phosphate (S3P) and
phosphoenolpyruvate to form 5-enolpyruvyl-shikimate-3-phosphate (EPSP). Glyphosate is ab-
sorbed through foliage and minimally through roots, meaning that it is only effective on actively
growing plants and cannot prevent seeds from germinating. After application, glyphosate is readi-
ly transported around the plant to growing roots and leaves and this systemic activity is important
for its effectiveness. Inhibiting the enzyme causes shikimate to accumulate in plant tissues and
diverts energy and resources away from other processes. While growth stops within hours of ap-
plication, it takes several days for the leaves to begin turning yellow.

EPSP is subsequently dephosphorylated to chorismate, an essential precursor for the amino acids
mentioned above. These amino acids are used in protein synthesis and to produce secondary me-
tabolites such as folates, ubiquinones, and naphthoquinone.

X-ray crystallographic studies of glyphosate and EPSPS show that glyphosate functions by occu-
pying the binding site of the phosphoenolpyruvate, mimicking an intermediate state of the ternary
enzyme substrates complex. Glyphosate inhibits the EPSPS enzymes of different species of plants
and microbes at different rates.

EPSPS is produced only by plants and microbes; the gene coding for it is not in the mammalian
genome.

Environmental Fate
Glyphosate adsorbs strongly to soil, and residues are expected to generally be immobile in soil.
Ground and surface water pollution is limited. Glyphosate is readily degraded by soil microbes to
aminomethylphosphonic acid (AMPA, which like glyphosate strongly adsorbs to soil solids and
is thus unlikely to leach to groundwater). Though both glyphosate and AMPA are commonly de-
tected in water bodies, a portion of the AMPA detected may actually be the result of degradation
of detergents rather than from glyphosate. Glyphosate does have the potential to contaminate
surface waters due to its aquatic use patterns and through erosion, as it adsorbs to soil particles
suspended in runoff. The mechanism of glyphosate sorption to soil is similar to that of phosphate
fertilizers, the presence of which can reduce glyphosate sorption. Phosphate fertilizers are subject

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to release from sediments into water bodies under anaerobic conditions, and similar release can
also occur with glyphosate, though significant impact of glyphosate release from sediments has
not been established. Limited leaching can occur after high rainfall after application. If glyphosate
reaches surface water, it is not broken down readily by water or sunlight.

The half-life of glyphosate in soil ranges between 2 and 197 days; a typical field half-life of 47 days
has been suggested. Soil and climate conditions affect glyphosate’s persistence in soil. The me-
dian half-life of glyphosate in water varies from a few to 91 days. At a site in Texas, half-life was
as little as three days. A site in Iowa had a half-life of 141 days. The glyphosate metabolite AMPA
has been found in Swedish forest soils up to two years after a glyphosate application. In this case,
the persistence of AMPA was attributed to the soil being frozen for most of the year. Glyphosate
adsorption to soil, and later release from soil, varies depending on the kind of soil. Glyphosate is
generally less persistent in water than in soil, with 12- to 60-day persistence observed in Canadian
ponds, although persistence of over a year has been recorded in the sediments of American ponds.
The half-life of glyphosate in water is between 12 days and 10 weeks.

According to the National Pesticide Information Center fact sheet, glyphosate is not included in
compounds tested for by the Food and Drug Administration’s Pesticide Residue Monitoring Pro-
gram, nor in the United States Department of Agriculture’s Pesticide Data Program. However, a
field test showed that lettuce, carrots, and barley contained glyphosate residues up to one year
after the soil was treated with 3.71 lb of glyphosate per acre (4.15 kg per hectare). The U.S. has de-
termined the acceptable daily intake of glyphosate at 1.75 milligrams per kilogram of bodyweight
per day (mg/kg/bw/day) while the European Union has set it at 0.3.

Use

Estimated use of glyphosate in the USA in 2013 and estimated total use from 1992–2013

Glyphosate is effective in killing a wide variety of plants, including grasses and broadleaf and
woody plants. By volume, it is one of the most widely used herbicides. In 2007, glyphosate was
the most used herbicide in the United States agricultural sector, with 180 to 185 million pounds

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252 Fundamentals of Weed Science

(82,000 to 84,000 tonnes) applied, the second-most used in home and garden with 5 to 8 million
pounds (2,300 to 3,600 tonnes) and government applied 13 to 15 million pounds (5,900 to 6,800
tonnes) in industry and commerce. It is commonly used for agriculture, horticulture, viticulture,
and silviculture purposes, as well as garden maintenance (including home use). It has a relatively
small effect on some clover species and morning glory.

Glyphosate used as an alternative to mowing in an apple orchard in Ciardes, Italy

Glyphosate and related herbicides are often used in invasive species eradication and habitat res-
toration, especially to enhance native plant establishment in prairie ecosystems. The controlled
application is usually combined with a selective herbicide and traditional methods of weed eradi-
cation such as mulching to achieve an optimal effect.

In many cities, glyphosate is sprayed along the sidewalks and streets, as well as crevices in between
pavement where weeds often grow. However, up to 24% of glyphosate applied to hard surfaces can
be run off by water. Glyphosate contamination of surface water is attributed to urban and agricul-
tural use. Glyphosate is used to clear railroad tracks and get rid of unwanted aquatic vegetation.
Since 1994, glyphosate has been used in aerial spraying in Colombia in coca eradication programs;
Colombia announced in May 2015 that by October, it would cease using glyphosate in these pro-
grams due to concerns about human toxicity of the chemical.

In addition to its use as an herbicide, glyphosate is also used for crop desiccation (siccation) to in-
crease harvest yield, and as a result of desiccation, to increase sucrose concentration in sugarcane
before harvest.

Genetically Modified Crops


Some micro-organisms have a version of 5-enolpyruvoyl-shikimate-3-phosphate synthetase (EP-
SPS) resistant to glyphosate inhibition. A version of the enzyme that was both resistant to glypho-
sate and that was still efficient enough to drive adequate plant growth was identified by Monsan-
to scientists after much trial and error in an Agrobacterium strain called CP4, which was found
surviving in a waste-fed column at a glyphosate production facility. This CP4 EPSPS gene was
cloned and transfected into soybeans. In 1996, genetically modified soybeans were made commer-

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Weed Control: Methods and Techniques 253

cially available. Current glyphosate-resistant crops include soy, maize (corn), canola, alfalfa, sugar
beets, and cotton, with wheat still under development.

In 2015, 89% of corn, 94% of soybeans, and 89% of cotton produced in the US were genetically
modified to be herbicide-tolerant.

Formulations and Tradenames


Glyphosate is marketed in the United States and worldwide by many agrochemical companies, in
different solution strengths and with various adjuvants, under dozens of tradenames. As of 2010,
more than 750 glyphosate products were on the market. In 2012, in terms of volume about half of
the total global consumption of glyphosate was for conventional crops; Asia Pacific was the largest
and fastest growing market. Chinese manufacturers collectively are the world’s largest producers
of glyphosate and its precursors and account for about 30% of global exports. Key manufacturers
include Anhui Huaxing Chemical Industry Company, BASF, Bayer CropScience, Dow AgroSci-
ences, DuPont, Jiangsu Good Harvest-Weien Agrochemical Company, Monsanto, Nantong Jiang-
shan Agrochemical & Chemicals Co., Nufarm Limited, SinoHarvest, Syngenta, and Zhejiang Xinan
Chemical Industrial Group Company.

Glyphosate is an acid molecule, so it is formulated as a salt for packaging and handling. Various
salt formulations include isopropylamine, diammonium, monoammonium, or potassium as the
counterion. Some brands include more than one salt. Some companies report their product as acid
equivalent (ae) of glyphosate acid, or some report it as active ingredient (ai) of glyphosate plus the
salt, and others report both. To compare performance of different formulations, knowledge of how
the products were formulated is needed. Given that different salts have different weights, the acid
equivalent is a more accurate method of expressing and comparing concentrations.

Adjuvant loading refers to the amount of adjuvant already added to the glyphosate product. Fully
loaded products contain all the necessary adjuvants, including surfactant; some contain no adju-
vant system, while other products contain only a limited amount of adjuvant (minimal or partial
loading) and additional surfactants must be added to the spray tank before application. As of 2000
(just before Monsanto’s patent on glyphosate expired), over 400 commercial adjuvants from over
34 different companies were available for use in commercial agriculture.

Products are supplied most commonly in formulations of 120, 240, 360, 480, and 680 g/l of active
ingredient. The most common formulation in agriculture is 360 g/l, either alone or with added
cationic surfactants.

For 360 g/l formulations, European regulations allow applications of up to 12 l/ha for control of
perennial weeds such as couch grass. More commonly, rates of 3 l/ha are practiced for control of
annual weeds between crops.

Monsanto
Monsanto developed and patented the use of glyphosate to kill weeds in the early 1970s, and first
brought it to market in 1974, under the Roundup brandname. While its initial patent expired in
1991, Monsanto retained exclusive rights in the United States until its patent on the isopropyl-
amine salt expired in September 2000.

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254 Fundamentals of Weed Science

Monsanto’s Roundup is the earliest formulation of glyphosate.

As of 2009, sales of these herbicide products represented about 10% of Monsanto’s revenue due to
competition from other producers of other glyphosate-based herbicides; their Roundup products
(which include GM seeds) represented about half of Monsanto’s gross margin.

The active ingredient of the Monsanto herbicides is the isopropylamine salt of glyphosate. Another
important ingredient in some formulations is the surfactant polyethoxylated tallow amine.

Monsanto also produces seeds which grow into plants genetically engineered to be tolerant to
glyphosate. The genes contained in these seeds are patented. Such crops allow farmers to use gly-
phosate as a postemergence herbicide against most broadleaf and cereal weeds. Soy was the first
glyphosate-resistant crop.

Toxicity
Glyphosate is the active ingredient in herbicide formulations containing it. However, in addition
to glyphosate salts, commercial formulations of glyphosate contain additives such as surfactants
which vary in nature and concentration. The surfactants are added to enable the glyphosate to
penetrate the cuticle of the plants. Toxicologists have studied glyphosate alone and formulations.

Glyphosate Alone
Humans
Many regulatory and scholarly reviews have evaluated the relative toxicity of glyphosate as an
herbicide. The German Federal Institute for Risk Assessment toxicology review in 2013 found
that “the available data is contradictory and far from being convincing” with regard to correlations
between exposure to glyphosate formulations and risk of various cancers, including non-Hodgkin
lymphoma.

Early epidemiological studies did not find associations between long-term, low-level exposure to
glyphosate and any disease. A 2000 review concluded that “under present and expected conditions
of new use, there is no potential for Roundup herbicide to pose a health risk to humans”. A 2002
review by the European Union reached the same conclusion. In 2013 the European commission
reviewed a 2002 finding that had concluded equivocal evidence existed of a relationship between

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Weed Control: Methods and Techniques 255

glyphosate exposure during pregnancy and cardiovascular malformations and found that “there is
no increased risk at the levels of exposure below those that caused maternal toxicity.” A 2013 re-
view found that neither glyphosate nor typical glyphosate-based formulations pose a genotoxicity
risk in humans under normal conditions of human or environmental exposures.

A 2014 review article reported a significant association between B-cell lymphoma and glypho-
sate occupational exposure. In March 2015, the World Health Organization’s International Agency
for Research on Cancer classified glyphosate as “probably carcinogenic in humans” (category 2A)
based on epidemiological studies, animal studies, and in vitro studies. However, in 2016 a joint
meeting of the United Nations (FAO) Panel of Experts on Pesticide Residues in Food and the
Environment and the World Health Organization Core Assessment Group on Pesticide Residues
concluded that based on the available evidence “glyphosate is unlikely to pose a carcinogenic risk
to humans from exposure through the diet”.

Other Mammals
Amongst mammals, glyphosate is considered to have “low to very low toxicity”. The LD50 of gly-
phosate is 5,000 mg/kg for rats, 10,000 mg/kg in mice and 3,530 mg/kg in goats. The acute der-
mal LD50 in rabbits is greater than 2,000  mg/kg. Indications of glyphosate toxicity in animals
typically appear within 30 to 120 minutes following ingestion of a large enough dose, and include
initial excitability and tachycardia, ataxia, depression, and bradycardia, although severe toxicity
can develop into collapse and convulsions.

A review of unpublished short-term rabbit-feeding studies reported severe toxicity effects at


150 mg/kg/day and “no observed adverse effect level” doses ranging from 50 to 200 mg/kg/day.

Glyphosate can have carcinogenic effects in nonhuman mammals. These include the induction of
positive trends in the incidence of renal tubule carcinoma and haemangiosarcoma in male mice,
and increased pancreatic islet-cell adenoma in male rats.

Glyphosate-based herbicides may cause life-threatening arrhythmias in mammals. Evidence also


shows that such herbicides cause direct electrophysiological changes in the cardiovascular systems
of rats and rabbits.

Other Aquatic Fauna


In freshwater invertebrates (species unspecified), glyphosate has a 48-hour LC50 ranging from 55 to
780 ppm. The 96-hour LC50 is 281 ppm for grass shrimp (Palaemonetas vulgaris) and 934 ppm for
fiddler crabs (Uca pagilator). These values make glyphosate “slightly toxic to practically non-toxic”.

Antimicrobial Activity
The antimicrobial activity of glyphosate has been described in the microbiology literature since its
discovery in 1970 and the description of glyphosate’s mechanism of action in 1972. Efficacy was
described for numerous bacteria and fungi. Glyphosate can control the growth of apicomplexan
parasites, such as Toxoplasma gondii, Plasmodium falciparum (malaria), and Cryptosporidium
parvum, and has been considered an antimicrobial agent in mammals. Inhibition can occur with
some Rhizobium species important for soybean nitrogen fixation, especially under moisture stress.

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256 Fundamentals of Weed Science

Soil Biota

Degradation pathway of glyphosate in the ground

When glyphosate comes into contact with the soil, it can be bound to soil particles, thereby slowing
its degradation. Glyphosate and its degradation product, aminomethylphosphonic acid, residues
are considered to be much more benign toxicologically and environmentally than most of the her-
bicides replaced by glyphosate.

Other studies found that while “recommended dosages of glyphosate did not affect growth rates”,
much higher dosages reduced respiration in nitrogen-fixing bacteria.

Glyphosate can be harmful to some earthworms such as Apporectodea caliginosa, even at very low
doses, and causes a significant reduction in growth of the earthworm Eisenia fetida.

Glyphosate-based Formulations
Glyphosate-based formulations may contain a number of adjuvants, the identities of which are
considered trade secrets. Surfactants are used in herbicide formulations as wetting agents, to max-
imize coverage and aid penetration of the herbicide(s) through plant leaves. As agricultural spray
adjuvants, surfactants may be mixed into commercial formulations, such as Roundup, or they may
be purchased separately and mixed on-site (tank mix).

Polyethoxylated tallow amine (POEA) is a surfactant used in the original Roundup formulation
and was still being commonly used in 2015. Different versions of Roundup have included different
percentages of POEA. Although Monsanto product fact sheets do not disclose surfactants and their
percentages, a 1997 US government report said that Roundup is 15% POEA while Roundup Pro is
14.5%. A review of the literature provided to the EPA in 1997 found that POEA was more toxic to
fish than glyphosate was. POEA is more toxic to fish and amphibians than glyphosate alone.

Human
A 2000 review concluded that “under present and expected conditions of new use, there is no po-
tential for Roundup herbicide to pose a health risk to humans”. A 2002 review by the European
Union reached the same conclusion.

A 2012 meta-analysis of all epidemiological studies of exposure to glyphosate formulations found


no correlation with any kind of cancer. The 2013 systematic review by the German Institute for
Risk Assessment of epidemiological studies of workers who use pesticides, exposed to glyphosate
formulations found no significant risk, stating that “the available data are contradictory and far

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Weed Control: Methods and Techniques 257

from being convincing”.:Volume 1, p64-66 However, a 2014 meta-analysis of the same studies found a
correlation between occupational exposure to glyphosate formulations and increased risk of B cell
lymphoma, the most common kind of non-Hodgkin lymphoma. Workers exposed to glyphosate
were about twice as likely to get B cell lymphoma.

Acute toxicity is dose-related; Skin exposure to ready-to-use glyphosate formulations can cause irrita-
tion, and photocontact dermatitis has been occasionally reported. These effects are probably due to the
preservative benzisothiazolin-3-one. Severe skin burns are very rare. Inhalation is a minor route of ex-
posure, but spray mist may cause oral or nasal discomfort, an unpleasant taste in the mouth, or tingling
and irritation in the throat. Eye exposure may lead to mild conjunctivitis. Superficial corneal injury is
possible if irrigation is delayed or inadequate. Death has been reported after deliberate overdose. In-
gestion of Roundup ranging from 85 to 200 ml (of 41% solution) has resulted in death within hours of
ingestion, although it has also been ingested in quantities as large as 500 ml with only mild or moderate
symptoms. Consumption of over 85 ml of concentrated product are likely to cause serious symptoms
in adults including burns due to corrosive effects as well as kidney and liver damage. More severe cas-
es cause “respiratory distress, impaired consciousness, pulmonary edema, infiltration on chest X-ray,
shock, arrhythmias, renal failure requiring haemodialysis, metabolic acidosis, and hyperkalaemia” and
death is often preceeded by bradycardia and ventricular arrhythmias.

Other Animals
A 2000 review of the ecotoxicological data on Roundup shows at least 58 studies exist on the ef-
fects of Roundup on a range of organisms. This review concluded, “...for terrestrial uses of Round-
up minimal acute and chronic risk was predicted for potentially exposed non-target organisms”.

In reproductive toxicity studies performed in rats and rabbits, no adverse maternal or offspring
effects were seen at doses below 175–293 mg/kg of body weight per day.

Fish
Monsanto and other companies produce glyphosate products with alternative surfactants specifical-
ly formulated for aquatic use, for example the Monsanto products “Biactive” and “AquaMaster”. Gly-
phosate formulations are much more toxic for amphibians and fish than glyphosate alone. The half-
life of POEA (21–42 days) is longer than that for glyphosate (7–14 days) in aquatic environments.

Amphibians
Some researchers have suggested the toxicity effects of pesticides on amphibians may be different from
those of other aquatic fauna because of their lifestyle; amphibians may be more susceptible to the toxic
effects of pesticides because they often prefer to breed in shallow, lentic, or ephemeral pools. These
habitats do not necessarily constitute formal water-bodies and can contain higher concentrations of
pesticide compared to larger water-bodies. Studies in a variety of amphibians have shown the toxicity
of GBFs containing POEA to amphibian larvae. These effects include interference with gill morphology
and mortality from either the loss of osmotic stability or asphyxiation. At sub-lethal concentrations,
exposure to POEA or glyphosate/POEA formulations have been reported to be associated with delayed
development, accelerated development, reduced size at metamorphosis, developmental malformations
of the tail, mouth, eye and head, histological indications of intersex and symptoms of oxidative stress.

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258 Fundamentals of Weed Science

A 2003 study of various formulations of glyphosate found, “[the] risk assessments based on esti-
mated and measured concentrations of glyphosate that would result from its use for the control
of undesirable plants in wetlands and over-water situations showed that the risk to aquatic or-
ganisms is negligible or small at application rates less than 4 kg/ha and only slightly greater at
application rates of 8 kg/ha.”

A 2013 meta-analysis reviewed the available data related to potential impacts of glyphosate-based her-
bicides on amphibians. According to the authors, the use of glyphosate-based pesticides cannot be con-
sidered the major cause of amphibian decline, the bulk of which occurred prior to the widespread use of
glyphosate or in pristine tropical areas with minimal glyphosate exposure. The authors recommended
further study of species- and development-stage chronic toxicity, of environmental glyphosate levels,
and ongoing analysis of data relevant to determining what if any role glyphosate might be playing in
worldwide amphibian decline, and suggest including amphibians in standardized test batteries.

Other Aquatic Fauna


Glyphosate-based formulations can cause oxidative stress in bullfrog tadpoles and Pacific oysters.

Effect on Plant Health


A correlation was found between an increase in the infection rate of wheat by Fusarium head blight
and the application of glyphosate, but “because of the nature of this study, we could not determine
if the association between previous GF (glyphosate formulation) use and FHB development was a
cause-effect relationship”. Other studies have found causal relationships between glyphosate and
decreased disease resistance. Exposure to glyphosate has been shown to change the species com-
position of endophytic bacteria in plant hosts, which is highly variable.

Endocrine Disruption
In 2007, the EPA selected glyphosate for further screening through its Endocrine Disruptor Screen-
ing Program (EDSP). Selection for this program is based on a compound’s prevalence of use and
does not imply particular suspicion of endocrine activity. On June 29, 2015 the EPA released Weight
of Evidence Conclusion of the EDSP Tier 1 screening for glyphosate, recommending that glyphosate
not be considered for Tier 2 testing. The Weight of Evidence conclusion stated “...there was no con-
vincing evidence of potential interaction with the estrogen, androgen or thyroid pathways.”

Genetic Damage
Several studies have not found mutagenic effects, so glyphosate has not been listed in the United
States Environmental Protection Agency or the International Agency for Research on Cancer data-
bases. Various other studies suggest glyphosate may be mutagenic. The IARC monograph noted that
glyphosate-based formulations can cause DNA strand breaks in various taxa of animals in vitro

Government and Organization Positions


European Food Safety Authority
A 2013 systematic review by the German Institute for Risk Assessment (BfR) examined more than

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Weed Control: Methods and Techniques 259

1000 epidemiological studies, animal studies, and in vitro studies. It found that “no classification
and labelling for carcinogenicity is warranted” and did not recommend a carcinogen classification
of either 1A or 1B. It provided the review to EFSA in January 2014 which published it in December
2014. On November, 12th, 2015, EFSA published its conclusion on the risk assessment of glypho-
sate, stating it was “unlikely to pose a carcinogenic hazard to humans”.

EFSA’s decision and the BfR report were criticized in an open letter published by 96 scientists in
November 2015 saying that the BfR report failed to adhere to accepted scientific principles of open
and transparent procedures. The BfR report included unpublished data, lacked authorship, omit-
ted references, and did not disclose conflict-of-interest information.

On April 4, 2016, Dr. Vytenis Andriukaitis, European Commissioner for Health and Food Safety,
wrote an open letter to the Chair of the Board of the Glyphosate Task at Monsanto Europe asking
to publish the full studies provided to the EFSA.

US Environmental Protection Agency


The EPA, which last reviewed glyphosate in 1993, considers glyphosate to be noncarcinogenic and
relatively low in dermal and oral acute toxicity. The EPA considered a “worst case” dietary risk
model of an individual eating a lifetime of food derived entirely from glyphosate-sprayed fields
with residues at their maximum levels. This model indicated that no adverse health effects would
be expected under such conditions. In 2015, the EPA initiated a review glyphosate’s toxicity and in
2016 reported their conclusion that glyphosate is likely not carcinogenic.

International Agency for Research on Cancer


In March 2015, the International Agency for Research on Cancer published a summary of their
forthcoming monograph on glyphosate, and classified glyphosate as “probably carcinogenic in hu-
mans” (category 2A) based on epidemiological studies, animal studies, and in vitro studies. It
noted that there was “limited evidence” of carcinogenicity in humans for non-Hodgkin lymphoma.
The IARC classifies substances for their carcinogenic potential, and “a few positive findings can be
enough to declare a hazard, even if there are negative studies, as well.” Unlike the BfR, it does not
conduct a so-called risk assessment weighing benefits against risk.

The BfR responded that IARC reviewed only a selection of what they had reviewed earlier, and ar-
gued that other studies, including a cohort study called ‘Agricultural Health Study’, do not support
the classification. The IARC report did not include the German regulatory study published in Decem-
ber 2014, nor did it include industry-funded studies. Monsanto called the IARC report biased and
said it wanted it to be retracted. It started a case against California’s carcinogen classification in 2016.

Effects of use
Emergence of Resistant Weeds
In the 1990s, when the first genetically modified crops-such as glyphosate-resistant corn, canola,
soybean and cotton—were introduced, no glyphosate-resistant weeds existed. By 2014, glypho-
sate-resistant weeds dominated herbicide-resistant research. At that time, 23 glyphosate-resistant
species were found in 18 countries.

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260 Fundamentals of Weed Science

“Resistance evolves after a weed population has been subjected to intense selection pressure in
the form of repeated use of a single herbicide.” Weeds resistant to the herbicide have been called
‘superweeds’.

According to Ian Heap, a weed specialist, who completed his PhD on resistance to multiple herbi-
cides in annual ryegrass (Lolium rigidum) in 1988 –the first case of an herbicide-resistant weed in
Australia–by 2014 the Lolium rigidum was the “world’s worst herbicide-resistant weed with instanc-
es in “12 countries, 11 sites of action, 9 cropping regimens” and affecting over 2 million hectares. An-
nual ryegrass was known to be resistant to herbicides since 1982. By 1996, the first documented case
of glyphosate-resistant L. rigidum was reported in Australia in 1996 near Orange, New South Wales.
In 2006, farmers associations were reporting 107 biotypes of weeds within 63 weed species with
herbicide resistance. In 2009, Canada identified its first resistant weed, giant ragweed, and at that
time 15 weed species had been confirmed as resistant to glyphosate. As of 2010, in the United States
7 to 10 million acres (2.8 to 4.0 million hectares) of soil were afflicted by superweeds, or about 5% of
the 170 million acres planted with corn, soybeans, and cotton, the crops most affected, in 22 states.
In 2012, Charles Benbrook reported that the Weed Science Society of America listed 22 superweeds
in the U.S., with over 5.7×10^6 ha (14×10^6 acres) infested by GR weeds and that Dow AgroSciences
had carried out a survey and reported a figure of around 40×10^6 ha (100×10^6 acres). The Inter-
national Survey of Herbicide Resistant Weeds database lists species that are resistant to glyphosate.

In response to resistant weeds, farmers are hand-weeding, using tractors to turn over soil between
crops, and using other herbicides in addition to glyphosate.

Monsanto scientists have found that some resistant weeds have as many as 160 extra copies of a
gene called EPSPS, the enzyme glyphosate disrupts.

Palmer Amaranth

Amaranthus palmeri

In 2004, a glyphosate-resistant variation of Amaranthus palmeri, commonly known as Palmer


amaranth, was found in Georgia and confirmed by a 2005 study. In 2005, resistance was also
found in North Carolina. Widespread use of Roundup Ready crops led to an unprecedented se-
lection pressure, and glyphosate resistance followed. The weed variation is now widespread in the
southeastern United States. Cases have also been reported in Texas and Virginia.

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Weed Control: Methods and Techniques 261

Conyza

Conyza canadensis

Conyza bonariensis (also known as hairy fleabane and buva) and Conyza canadensis (known as
horseweed or marestail), are other weed species that had lately developed glyphosate resistance.
A 2008 study on the current situation of glyphosate resistance in South America concluded “resis-
tance evolution followed intense glyphosate use” and the use of glyphosate-resistant soybean crops
is a factor encouraging increases in glyphosate use. In the 2015 growing season, glyphosate-resis-
tant marestail proved to be especially problematic to control in Nebraska production fields.

Ryegrass

Ryegrass Lolium perenne

Glyphosate-resistant ryegrass (Lolium) has occurred in most of the Australian agricultural areas
and other areas of the world. All cases of evolution of resistance to glyphosate in Australia were
characterized by intensive use of the herbicide while no other effective weed control practices were
used. Studies indicate the resistant ryegrass does not compete well against nonresistant plants and
their numbers decrease when not grown under conditions of glyphosate application.

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262 Fundamentals of Weed Science

Johnson Grass
Glyphosate-resistant Johnson grass (Sorghum halepense) is found in glyphosate-resistant soy-
bean cultivation in northern Argentina.

Monarch Butterfly
Use of glyphosate to clear milkweed along roads and fields may have contributed to a decline
in monarch butterfly populations in the Midwest. Along with deforestation and adverse weather
conditions, the decrease in milkweed contributed to an 81% decline in monarchs. The Natural Re-
sources Defense Council (NRDC) filed a suit in 2015 against the EPA, in which it is argued that the
agency ignored warnings about the dangers of glyphosate usage for monarchs.

Legal Status
Glyphosate was first approved for use in the 1970s, and as of 2010 was labelled for use in 130
countries.

In September 2013, the legislative assembly of El Salvador approved legislation to ban 53 agro-
chemicals, including glyphosate; the ban on glyphosate was set to begin in 2015.

In April 2014, the legislature of the Netherlands passed legislation prohibiting sale of glyphosate
to individuals for use at home; commercial sales were not affected.

In May 2015, the president of Sri Lanka banned the use and import of glyphosate, effective imme-
diately.

In May 2015, Bermuda blocked importation on all new orders of glyphosate-based herbicides for
a temporary suspension awaiting outcomes of research.

In May 2015, Colombia announced that it would stop using glyphosate by October 2015 in the
destruction of illegal plantations of coca, the raw ingredient for cocaine. Farmers have complained
that the aerial fumigation has destroyed entire fields of coffee and other legal produce.

In June 2015, the French Ecology Minister asked nurseries and garden centers to halt over-the-
counter sales of glyphosate in the form of Monsanto’s Roundup. This was a nonbinding request
and all sales of glyphosate remain legal in France until 2022, when the substance will be banned
for home gardening.

A vote on the relicencing of glyphosate in the EU stalled in March 2016. Member states France,
Sweden, and the Netherlands objected to the renewal. A vote to reauthorize on a temporary ba-
sis failed in June 2016 but at the last-minute the license was extended for 18 months and will be
re-evaluated at the end of 2017.

Legal Cases
Advertising Controversy
The New York Times reported that in 1996, “Dennis C. Vacco, the Attorney General of New York,

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Weed Control: Methods and Techniques 263

ordered the company Monsanto to pull ads that said Roundup was “safer than table salt” and
“practically nontoxic” to mammals, birds and fish. The company withdrew the spots, but also said
that the phrase in question was permissible under E.P.A. guidelines.”

In 2001, French environmental and consumer rights campaigners brought a case against Mon-
santo for misleading the public about the environmental impact of its herbicide Roundup, on the
basis that glyphosate, Roundup’s main component, is classed as “dangerous for the environment”
and “toxic for aquatic organisms” by the European Union. Monsanto’s advertising for Roundup
had presented it as biodegradable and as leaving the soil clean after use. In 2007, Monsanto was
convicted of false advertising and was fined 15,000 euros. Monsanto’s French distributor Scotts
France was also fined 15,000 euros. Both defendants were ordered to pay damages of 5,000 euros
to the Brittany Water and Rivers Association and 3,000 euros to the Consommation Logement
Cadre de vie, one of the two main general consumer associations in France. Monsanto appealed
and the court upheld the verdict; Monsanto appealed again to the French Supreme Court, and in
2009 it also upheld the verdict.

Trade Dumping Allegations


United States companies have cited trade issues with glyphosate being dumped into the western
world market areas by Chinese companies and a formal dispute was filed in 2010.

References
• Bleasdale, J. K. A.; Salter, Peter John (1 January 1991). The Complete Know and Grow Vegetables. Oxford Uni-
versity Press. ISBN 978-0-19-286114-6.

• Ross, Merrill A.; Lembi, Carole A. (2008). Applied Weed Science: Including the Ecology and Management of
Invasive Plants. Prentice Hall. p. 123. ISBN 978-0135028148.

• National Research Council (2002). Making Aquatic Weeds Useful: Some Perspectives for Developing Coun-
tries. The Minerva Group. pp. 1–9. ISBN 978-0-89499-180-6.

• National Research Council (2002). Making Aquatic Weeds Useful: Some Perspectives for Developing Coun-
tries. The Minerva Group. pp. 67–68. ISBN 978-0-89499-180-6.

• R. Goyal, Megh (2012). Management of drip/trickle or micro irrigation. Oakville, CA: Apple Academic Press.
p. 104. ISBN 978-1926895123.

• Alfred J. Turgeon; Lambert Blanchard McCarty; Nick Edward Christians (2009). Weed control in turf and
ornamentals. Prentice Hall. p. 126. ISBN 978-0-13-159122-6.

• Mahesh K. Upadhyaya; Robert E. Blackshaw (2007). Non-chemical Weed Management: Principles, Concepts
and Technology. CABI. pp. 135–. ISBN 978-1-84593-291-6.

• Stout, Ruth. Gardening Without Work. Devon-Adair Press, 1961. Reprinted by Norton Creek Press, 2011, pp.
6-7. ISBN 978-0-9819284-6-3

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6
Laws Related to Weed
The Weeds Act 1959 is an act that is regarding the control of several injurious weed species. This
act is restricted to the United Kingdom. International Plant Protection Convention and the Federal
Noxious Weed Act of 1974 are some of the laws related to weed control. This chapter discusses the
laws related to weeds in a critical manner providing key analysis to the subject matter.

Weeds Act 1959


The Weeds Act 1959 (7 & 8 Eliz. II c. 54) is an Act of the Parliament of the United Kingdom regard-
ing the control of several injurious weed species throughout the UK. It received Royal Assent on 16
July 1959, and aims to prevent the spread of the Broad Leaved Dock, Common Ragwort, Creeping
Thistle, Curled Dock and the Spear Thistle. It allows the Secretary of State, or any person acting on
their behalf or the Secretary of State for Environment, Food and Rural Affairs, to use measures of
enforcement to prevent the spread of weeds on private land, which, if not adhered to by the owner
of said land, can lead to a fine up to £1000 and further punishment.

In Scotland, powers are now exercisable by the Scottish Ministers rather than the Secretary
of State. This Act is amended for England and Wales by the Ragwort Control Act 2003. From
2014, the Anti-Social Behaviour, Crime and Policing Act allowed a community protection no-
tice to be issued for a range of nuisances, including an unreasonable failure to act over an
issue such as weed control, of any species. Ignoring such a notice could then lead to an ASBO
or criminal fine.

Species Referred to in the Weeds Act 1959


Broad Leaved Dock
The Broad Leaved Dock is a resilient and common perennial plant found in grasslands through-
out the United Kingdom. Unaffected by regular climatic variations and all but the most acidic
soils, the Broad Leaved Dock can produce around 60,000 seeds a year and flowers from June
to October. The seeds can survive for up to 50 years in soil due to a ‘chemical that inhibits
microbial decay’.

Curled Dock
The Curled Dock is found mostly in meadowland, wasteland, sand dunes, and dry soils. The Curled
Dock can be annual, biennial, or perennial, and can produce 3000 to 4000 seeds per plant. Al-
though said to contribute to animal wellbeing by providing nutrients which would otherwise be
absent, it also contains high levels of oxalic acid which could be damaging to stock.

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Laws Related to Weed 265

Creeping Thistle
The Creeping Thistle is a common resilient perennial plant found in grasslands throughout the
United Kingdom. Characterised by spined lobed leaves, it stands up to one metre high and blooms
with light purple flowers between July and September. Its root system is very deep, extending up
to three metres underground and six metres sideways; as such, it is very hard to remove from an
affected area. It competes fiercely with other plants or crops and can release a natural biocide into
the soil to inhibit growth of other species.

Ragwort
The Common Ragwort is a biennial yellow angiosperm which can grow to 30–100 cm high. Prolific
in seed distribution, a ragwort plant can produce up at 30,000 to 120,000 seeds. Toxic to cattle,
horses, pigs, deer, and goats, the ragwort has a high alkaloid concentration which causes liver
damage. Sheep are less affected but can suffer from consumption as the effect is ‘cumulative’.

Spear Thistle
The Spear Thistle is an annual or biennial plant which forms dark purple or reddish flowers above
dark green spiked leaves. A plant found in pastoral land and along roads, it is easily spread by ve-
hicles as they pass by. A severely competitive plant, it can eliminate pastoral crop and open crops
to infestation by insects; the spikes can dislodge from the plant and attach to wool, presenting a
problem for shearers.

International Plant Protection Convention


The International Plant Protection Convention (IPPC) is a 1951 multilateral treaty overseen by the
Food and Agriculture Organization that aims to secure coordinated, effective action to prevent and
to control the introduction and spread of pests of plants and plant products. The Convention ex-
tends beyond the protection of cultivated plants to the protection of natural flora and plant prod-
ucts. It also takes into consideration both direct and indirect damage by pests, so it includes weeds.

The Convention created a governing body consisting of each party, known as the Commission on
Phytosanitary Measures, which oversees the implementation of the Convention. As of September
2015, the Convention has 182 parties, which includes 179 United Nations member states, the Cook
Islands, Niue, and the European Union. The Convention is recognized by the World Trade Orga-
nization’s (WTO) Agreement on the Application of Sanitary and Phytosanitary Measures (the SPS
Agreement) as the only international standard setting body for plant health.

While the IPPC’s primary focus is on plants and plant products moving in international trade,
the Convention also covers research materials, biological control organisms, germplasm banks,
containment facilities, food aid, emergency aid and anything else that can act as a vector for the
spread of plant pests – for example, containers, packaging materials, soil, vehicles, vessels and
machinery.

The IPPC was created by member countries of the Food and Agriculture Organization of the United

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266 Fundamentals of Weed Science

Nations. The IPPC places emphasis on three core areas: international standard setting, information
exchange and capacity development for the implementation of the IPPC and associated international
phytosanitary standards. The Secretariat of the IPPC is housed at FAO headquarters in Rome, Italy,
and is responsible for the coordination of core activities under the IPPC work program.

In recent years the Commission of Phytosanitary Measures of the IPPC has developed a strategic
framework with the objectives of:

• protecting sustainable agriculture and enhancing global food security through the preven-
tion of pest spread;

• protecting the environment, forests and biodiversity from plant pests;

• facilitating economic and trade development through the promotion of harmonized scien-
tifically based phytosanitary measures, and:

• developing phytosanitary capacity for members to accomplish the preceding three objec-
tives.

By focusing the Convention’s efforts on these objectives, the Commission on Phytosanitary Mea-
sures of the IPPC intends to:

• Protect farmers from economically devastating pest and disease outbreaks.

• Protect the environment from the loss of species diversity.

• Protect ecosystems from the loss of viability and function as a result of pest invasions.

• Protect industries and consumers from the costs of pest control or eradication.

• Facilitate trade through International Standards that regulate the safe movements of plants
and plant products.

• Protect livelihoods and food security by preventing the entry and spread of new pests of
plants into a country.

Federal Noxious Weed Act of 1974


The Federal Noxious Weed Act of 1974 (“FNWA”, Pub.L. 93–629, 88 Stat. 2148, enacted January
3, 1975) established a federal program to control the spread of noxious weeds. The United States
Secretary of Agriculture was given the authority to declare plants “noxious weeds”, and limit the
interstate spread of such plants without a permit. The Secretary has the authority to inspect, seize,
and destroy products, and quarantine areas, if necessary to contain, or limit the spread of such
weeds.

Amended in 1990
The Act was amended by the 1990 Farm Bill on November 28, 1990. The amendment requires that

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Laws Related to Weed 267

each federal land-managing agency, including the Bureau of Land Management, the National Park
Service, the U.S. Fish and Wildlife Service, and the U.S. Forest Service, do the following:

• Designate an office or person adequately trained in managing undesirable plant species to


develop and coordinate a program to control such plants on the agency’s land;

• Establish and adequately fund this plant management program through the agency’s bud-
get process;

• Complete and implement cooperative agreements (requirements for which are provided)
with the States regarding undesirable plants on agency land; and

• Establish integrated management systems (as defined in the Act) to control or contain un-
desirable plants targeted under the cooperative agreements.

Superseded in 2000
The Act was superseded in 2000 by the Plant Protection Act (Pub.L. 106–224) on June 20, 2000,
except for the introductory section of the FNWA, and the amendment of 1990 (section 15, 7 U.S.C.
§ 2801 note and 7 U.S.C. § 2814).

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________________________ WORLD TECHNOLOGIES ________________________


Index
A E
Abrus Precatorius, 8, 11-15 Edibility, 40, 42-43, 170, 174
Alkaloids, 54, 91-92, 95-97, 100-101, 109, 146-147 Edible, 4-5, 7, 19-20, 30, 35, 37, 39-40, 43, 45, 47, 54-55,
Allium Vineale, 47 70, 77, 90, 137, 141-142, 144, 154-155, 165-166, 172, 174,
186
Alternative Herbicides, 242-243
Amaranth, 5, 7, 150-159, 186, 260
F
Anaerobic (sour) Mulch, 225 False Dandelions, 168
Animal Feed, 31, 43, 88 Federal Noxious Weed Act Of 1974, 11, 264, 266
Aquatic Weed Harvester, 187, 194-197 First Herbicides, 233
Archaeological Use, 232 Food For Wildlife, 171
Arctium Lappa, 46-48
As A Noxious Weed, 162, 172 G
Garden Cultivators, 218
B Gastronomy, 19
Bark Chips, 220, 224-225 Goetzeoideae, 100
Biochemistry Of Resistance, 241 Grass Clippings, 222-223
Biodegradable Mulch, 227 Groundcovers (living Mulches), 226
Bioherbicide, 211-212
Blackberry, 45, 70-76, 165-166, 214 H
Brassicaceae, 80-81 Hedera Helix, 24, 26
Bromus Tectorum, 8, 16-17, 114, 118 Herbicide Group (labeling), 237
Bush Regeneration, 187, 212-214 Herbicide Rotations, 243
Herbicides, 1, 7, 10, 12, 30, 35, 47, 76, 133, 140-141, 143,
C 156-157, 180, 183, 187, 192-194, 196, 210-211, 216, 226
Cardoon, 8, 18-21 227, 232-244, 246-247, 251-252, 254-256, 258, 260, 262
Centaurea Cyanus, 76 Hoe (tool), 227
Cestroideae (browallioideae), 98
I
Clinical Trials, 177
International Plant Protection Convention, 264-265
Colored Mulch, 225
Invasive Species, 18, 21, 24, 27, 33, 53, 111-113, 115-125,
Commercial Cultivation, 73 127-128, 130, 132, 134, 136-141, 146, 148, 155, 165, 196,
Cowpea, 46, 88-91, 185 213-214, 252
Cultivars, 19-20, 27, 71, 74, 76, 169, 175, 183
Cultivation, 1, 7, 18-20, 24, 26, 30, 35, 45, 49, 53, 73-74, J
76, 90, 96, 125, 136-137, 145-146, 148, 154, 172, 179, 187, Juncus Effusus, 8, 22-24
202, 204, 217, 242, 245, 262
Cultivators, 1, 187, 215-219 K
Kudzu, 6, 8, 28-35, 110, 112, 119, 127, 165
D
Dangers, 171, 262 L
List Of Beneficial Weeds, 40
Depth Steaming With Vacuum, 199
Digitaria Ciliaris, 8, 21
M
Draw Hoes, 228-229 Medicinal Use, 46, 174, 177
Drip Irrigation, 187, 191, 205-211, 221, 227 Medicinal Uses, 23, 92, 171

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270 Index

Mimosa Pudica, 40, 50-54 Sheet Steaming, 198-201


Mulch, 1, 4, 38-40, 187, 190, 220-227 Shelter Plants, 45
Mulching (composting) Over Unwanted Plants, 226 Silybum Marianum, 150, 175-177
Soil Decontamination, 203
N Soil Disinfestation, 203
Nicotianoideae, 92, 99-100, 102
Soil Improvement, 31
Nitrogen Fixation, 36, 53, 179, 226, 255
Soil Solarization, 191, 202-204
Nutrient Content Of Seeds, 72
Soil Steam Sterilization, 143, 187, 191, 197, 202
Solanaceae, 91-92, 94-96, 98, 101, 108
O
On-site Mulch Production, 226 Solanoideae, 92, 95, 100, 103
Organic Herbicides, 236, 246 Stale Seed Bed, 187, 191, 204
Organic Mulches, 222, 225 Steaming With Hoods, 200
Ornamentals, 91, 109, 128, 156, 175, 204, 263 Steaming With Superheated Steam, 198
Synthetic Herbicides, 233, 244, 246
P
Partially Integrated Sandwich Steaming, 198, 201 T
Peat Moss, 222-223 Taraxacum, 4, 46, 166-171
Pest-repellent, 45 Taxonomy, 30, 50, 70, 80, 89, 98, 123, 131, 147, 150-151
Petunioideae, 92, 100 Toxicodendron Radicans, 150, 159-162
Plant Movement, 52 Trap Crops, 37, 45, 183
Plantago Major, 172-174 Treatment Of Poison Ivy Rash, 164
Polyethylene Mulch, 226 Trifolium Repens, 178, 180-181
Polypropylene Mulch, 226
W
Weed Control, 1, 6-7, 9-10, 24, 137, 157, 187, 189-193,
R 195, 197, 199, 201, 203-205, 207, 209, 211-213, 215-219,
Row Crop Cultivators, 217
221, 223, 225, 227-229, 231, 233-235, 237, 239-243, 245
Rumex, 10, 46, 54-70 247, 249, 251, 253, 255, 257, 259, 261, 263-264
Weed Harvesting Equipment, 195
S Weed Propagation, 188
Schizanthoideae, 95, 101
Weeds Act, 1959, 10
Schwenckioideae, 101
Wood Chips, 223, 225
Scuffle Hoes, 228-230
Woodchip Mulch, 223-224
Seed Dispersal, 136-138, 167

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