Jean-Marc Dufour-Dror (Ph.

D)
Invasive
in Israel
Plants
Alien

Alien Invasive Plants in Israel Jean-Marc Dufour-Dror (Ph.D)

Alien Invasive Plants in Israel
 Alien
Invasive
Plants
‫העמותה לשמירת טבע‬
(‫במזה״ת )ע״ר‬

in Israel
Jean-Marc Dufour-Dror (Ph.D)
Proposed by the late Aharon Har-Even

Scientific Editors
Dr. Ori Fragman-Sapir, Head Scientist, the Jerusalem Botanical Gardens
Dr. Margareta Walczak, chief botanist at the Israel Nature and Parks Authority
Dr. Tuvia Yaacoby, chief herbologist at the Plant Protection Services, Ministry of
Agriculture and Rural Development, Bet Dagan
Hagar Leshner, Collection Manager, the National Herbarium, the Hebrew
University, Jerusalem

English Language Editor

Dr. Esther Lachman

Graphic Editing
Yuval Tal Ltd. Jerusalem

Publishing
The Middle East Nature Conservation Promotion Association

Printing
Ahva, Jerusalem

All rights reserved.

No part of this document may be copied or used for any purpose without written
permission from the publisher.

2012

ISBN 978-965-90466-7-6
�
Contents

Acknowledgements 10
Foreword To The English Edition 11
Introduction 12
1 Invasive Plant Species: Basic Concepts
and General Information 15
1.1 Alien species, invasive species and invasion stages 15
1.2 A glimpse at the enemy release hypothesis 18
1.3 Invasive plants: general biological and ecological characteristics 20
1.4 The dynamics of invasive plants: Basic aspects and
examples from Israel’s alien taxa 26
1.5 Examples of lists of invasive plants from Mediterranean and
arid regions in the world 29

2 Invasive Plants In Natural Areas In Israel: A Review 33

2.1 The factors underlying the invasion of alien plant in
natural areas in Israel 33
2.1.1 Anthropogenic activities that introduced invasive plant
species in Israel 33
2.1.2 Factors that increased the invasibility of Israel’
natural ecosystems in recent decades 39

2.2 The major effects of invasive plants on natural ecosystems in Israel 42

2.2.1 Displacement of local plant species 42
2.2.2 Invasive transformers 44
2.2.3 The habitats most vulnerable to invasion by
alien plants in Israel 46

2.3 Managing invasive plants in Israel 52

2.3.1 Physical and chemical control methods applied in Israel:
an overview 54
2.3.2 The issue of biological control against invasive
plants in Israel 60
2.3.3 Prevention of further plant invasions in Israel 62

2.4 Selection of species included in the book 63

3 – Alien Invasive Plants Of Israel: Factsheets 67

Acacia cyclops A.Cunn ex G.Don Western Coastal Wattle 68
Acacia karroo Hayne Karroothorn 70
Acacia paradoxa DC. Kangaroo Thorn, Prickly Wattle 72
Acacia salicina Lindl. Willow Wattle, Broughton Willow 74
Acacia saligna (Labill.) H.L.Wendl. Golden Wreath Wattle, Blue-leafed Wattle76
Acacia victoriae Benth. Elegant Wattle, Bramble Wattle 80
Ailanthus altissima (Mill.) Swingle Tree-of-Heaven, Chineese Sumac 82
Ambrosia confertiflora DC Burr Ragweed, Slimleaf Bursage 84
Atriplex holocarpa F. Muell. Pop Saltbush 88
Azolla filiculoides Lam. Red Water Fern, Large Mosquito Fern 90
Carpobrotus edulis (L.) N.E. Br. Hottentot Fig, Iceplant 92
Conyza bonariensis (L.) Cronquist Flax Leaved Fleabane 94
Conyza albida Willd. Ex Spreng. Fleabane 94
Conyza canadensis (L.) Cronquist Canada Horseweed 98
Cyperus involucratus Rottb. Umbrella sedge 100
Cyperus odoratus L. Fragrant Flatsedge 102
Datura stramonium L. Common thornapple 104
Dodonaea viscosa L.(Jacq.) Hopbush, Varnish Tree 106
Eichhornia crassipes (Mart.) Solms Water Hyacinth 108
Eucalyptus camaldulensis Dehn. Murray Red Gum, River Redgum 112
Ficus benghalensis L. East Indian Figtree, Indian Banyan 114
Ficus microcarpa L.fil. Chinese Banyan, Curtain Fig 116
Ficus religiosa L. Bo Tree, Sacred Fig 118
Heterotheca subaxillaris (Lam.) Britton & Rusby Camphorweed 120
Ipomoea aquatica Forssk. Chinese Waterspinach, Water Spinach 124
Lantana camara L. Common Lantana, Largeleaf Lantana 126
Melia azedarach L. Chinaberry tree 130
Myriophyllum aquaticum (Vell.) Verdc. Parrot’s Feather 132
Nicotiana glauca Graham Tree Tobacco 134
Oenothera drummondii Hooker Beach Evening Primrose 136
Oxalis pes-caprae L. Bermuda Buttercup 138
Parkinsonia aculeata L. Horse Bean, Jerusalem Thorn 140
Paspalum distichum L. Couch Paspalum, Knotgrass 142
Pennisetum clandestinum Hochst. ex Chiov. Kikuyu Grass 144
Phytolacca americana L. American pokeweed, Pigeon berry 146
Pinus brutia Ten. Calabrian pine, Turkish red pine 150
Pistia stratiotes L. Water Lettuce, Tropical Duckweed 122
Populus alba L. Silver-Leaf Poplar, White Poplar 154
Prosopis juliflora (Sw.) DC. Mesquite, Devil’s Tree 156
Ricinus communis L. Castor Bean, Castor Oil Plant 158
Robinia pseudoacacia L. Black Locust, False Acacia 160
Salvinia molesta D.S.Mitch. Africa Payal, Giant Salvinia 162
Schinus molle L. Peruvian Pepper Tree 164
Schinus terebinthifolius Raddi. Brazilian Pepper Tree 166
Sesbania sesban (L.) Merr. Common sesban, Egyptian pea 168
Solanum elaeagnifolium Cav. Silverleaf Nightshade 170
Verbesina encelioides (Cav.) A.Gray Crown Beard, American Dogweed 172
Washingtonia robusta H. A. Wendl. Mexican Fan Palm 174
Washingtonia filifera (Lindl. ex Andre) H. A. California Fan Palm 174
Xanthium strumarium L. Common cocklebur, Rough cocklebur 176

Glossary 179
References 182
Appendix 203
 Conserving Biodiversity

Conserving biodiversity and ecosystem services is high on the priority list

of the global agenda. The United Nations declared 2010 the International
Year of Biodiversity and Israel’s Ministry of Environmental Protection,
together with the Israel Nature and Parks Authority and Israeli scientists
prepared a National Biodiversity Plan for Israel.

Biological invasions are considered the second most important factor,

after habitat destruction, contributing to the decrease of biodiversity in
the world. During the last decade, Israel too has had to deal with the
8 problem in natural and other open spaces, including protected areas.

“Alien Invasive Plants in Israel” presents the exotic species that have
become invasive in natural areas in Israel. Among invasive organisms,
plants are considered the “silent invaders” that displace local species
from their habitats, even in protected areas. Moreover, the spread of
invasive plants also leads to a gradual change in the character of affected
ecosystems, and eventually to a reduction in species diversity in affected
areas.

For a number of years now, the Ministry of Environmental Protection along

with the Nature and National Parks Authority has supported research,
surveys and development of treatment and management methods to deal
with the phenomenon of invasive plants in Israel. As a result, extensive
knowledge has accumulated related to these plants, which has been
collected in the book “Alien Invasive Plants in Israel”. We hope that the
publication of this book will increase awareness to the threat of invasive
species in Israel among the authorities and bodies dealing with the
subject as well as among the public at large.

Alona Sheafer-Karo
Director General,
Israel Ministry of Environmental Protection
Invasive Plants in Israel – A Serious Threat

This publication is unique compared to the material usually published by

the Israel Nature and Parks Authority. Normally, when we publish a guide
or any other book relating to species, we focus on means of conserving
or protecting the species, in its natural habitat or at times with the help
of non-natural methods. This book focuses on plant species we would
prefer not to find in nature in Israel, species that we work to remove from
nature. These are known as alien invasive species.

Invasive species are considered today to be one of three major factors

leading to species extinction on the face of the earth. Human beings move 9
species from place to place on the face of the globe. Species translocation
is occasionally deliberate, and sometimes it occurs randomly or due to
lack of attention. These species have a destructive effect on natural
habitats and the native species inhabiting them. They alter the character
of the habitats, and replace indigenous species. Thus, the native flora and
fauna are replaced by alien species whose original habitats are in other
countries.

The Israel Nature and Parks Authority invests intensive efforts in dealing
with these species. The most important response is preventing them from
entering the country, which requires the involvement and cooperation of
various authorities. Invasive species that are already established must
be identified and located and their effect on nature in Israel studied.
Management will be determined by the degree of damage they have
inflicted and by the actions it is possible to implement. The optimal result
is the complete removal of invasive species, but if this cannot be achieved
their populations must be reduced, or at the very least stopped from
spreading.

This book is a reflection of the intensive efforts invested these last years
in identifying alien invasive plant species in Israel, mapping their major
locations and defining means to prioritize management options for dealing
with problem.

Shaul Goldstein
Director General,
Israel Nature and Parks Authority
 � Acknowledgements
Special thanks to the four scientific editors of this book, Ori Fragman-Sapir, head
scientist of the Jerusalem Botanical Gardens, Margareta Walczak, chief botanist at
the Israel Nature and Parks Authority, and Tuvia Yaacoby chief herbologist at the
Plant Protection Services, Ministry of Agriculture and Rural Development, and Hagar
Leshner from the National Herbarium (the Hebrew University of Jerusalem) whose
expert comments contributed greatly to this book.

I would like to express my thanks to the people listed below whose help was very
valuable:
10 Dan Perry, the CRB Foundation
Esther Lachman, language editor for the English edition
Haim Kigel, the Faculty of Agriculture, Food and Environment,
the Hebrew University
Meir Forti, Yosef Karasso, Yan Landau, Zvi Mendel, Avi Perevolotsky,
Noam Seligman, Agricultural Research Organization (ARO), Volcani Center
Hava Goldstein, Ben Inbar, Didi Kaplan, Ori Liniel, Yoram Malka,
Talia Oron, Amos Sabah, Yehoshua Shkedy, Yiftach Sinai, Israel Nature
and Parks Authority
Menachem Zalutsky, Noa Steiner, Mona Farkas Berdugo, the Israel Ministry of
Environmental Protection
Bruce Maslin, Western Australia Herbarium, Department of Environment
and Conservation, Western Australia
Gordon C. Tucker, Herbarium Curator, Dept. of Biological Sciences,
Eastern Illinois University, USA
Wayne O’Sullivan, Department of Environment and Conservation,
Bentley, Western Australia
Fiona Impson, Plant Protection Research Institute, Stellenbosch, South Africa
Genevieve Thompson – Centre for Invasion Biology, Stellenbosch University,
South Africa
Liat Dufour-Dror
Ron Frumkin
�
Foreword to the English edition
This book was first written and edited in Hebrew as it was initially intended
for the Israeli public. Yet, since invasive plants essentially ignore regional and
political borders, I decided to publish an English edition so the information
displayed in this book can be shared with a larger public, first and foremost
from neighboring countries. Many of the alien taxa invading natural and semi-
natural habitats in the various regions of Israel may pose a threat for natural
areas in the surrounding countries in the eastern Mediterranean and in the
Near East as well. Moreover, this English edition may also be of some interest
for ecologists and environmentalists who seek information about invasive
plants in Mediterranean-type, semiarid and arid regions. 11

Since the issue of alien invasive plants in Israel is very dynamic the information
displayed for several species was updated, particularly regarding the extent
of their proliferation in Israel. Similarly, the scientific literature cited in the
English edition was also updated and includes several recent publications that
came out between late 2010 and early 2012.

Jerusalem, March 2012

J.M.D.D
 �
Introduction
The aim of this book is to present basic, but current information on the topic
of alien invasive plant species in Israel specifically, and to provide detailed up
to date information on the main invasive plant taxa found in natural areas
throughout Israel. The information is presented from an ecological standpoint,
and focuses on general traits of alien plants, their stages of invasion, the causes
underlying invasion of alien plant species in Israel, the consequences for local
ecosystems and what is done in Israel to battle invasive plant species. The book
also aims at raising awareness to the issue of invasive alien species in Israel,
12 particularly in natural and protected areas.

The book is divided into three chapters:

The first chapter is dedicated to the definitions of several basic concepts required
for understanding the topic of invasive plant ecology. An overview of the main
ecological and biological characteristics typical of invasive plants is presented in
the following section of this chapter, with all examples taken from invading taxa
in Israel. During the writing of the book we refer to several regional databases of
invasive plant species from other countries with Mediterranean and arid climates
similar to Israel’s. The reference to these regional databases and lists of invasive
alien plant species was necessary in order to verify the geographical extent
of several invasive taxa outside Israel. A brief review of these data sources is
presented at the end of the first chapter.

The second chapter presents a brief analysis of the major characteristics of

the phenomenon of invasive plants in natural areas in Israel. The first section
of this chapter discusses the factors underlying the spread of invasive plants
in Israel. The second section reviews their main effects on local ecosystems,
focusing on habitats that are particularly susceptible to invasion. In the third
section we present a summary of the control techniques and methods currently
implemented in Israel to cope with the proliferation of invasive plants in
natural areas. The final section discusses the criteria for choosing the 50 major
invasive plant taxa in Israel whose characteristics are specified in the following
chapter, and how the ecological threat posed by each invasive species for Israeli
ecosystems was calculated.

The third chapter that makes up the bulk of this book provides detailed and up
to date information on the 50 most significant species of alien plants that invade
natural and semi-natural habitats in Israel. The information includes a short
description of the plant, information on its natural range (origin), background
related to its introduction to Israel, information on its known biological and
ecological traits, a description of its invasive status in Israel and an explanation
of how it affects local ecosystems. Existing and relevant control methods are
described for each species. The main regions in the world invaded by the species
are mentioned, and a brief list of references is presented for each species.

Finally there is an appendix with a current list of all the alien plant species
recorded in natural and disturbed areas in Israel up to early 2010. This list,
prepared by the author with Dr. Ori Fragman-Sapir from the Jerusalem Botanical
Gardens is the first in Israel since the list published by Dafni & Heller twenty
years ago.

Target audience and general scope of this book 13

This book is intended for ecologists, environmental consultants, landscape
architects, planners, managers of natural areas, students and anyone interested
in the phenomenon of invasive plants in Israel in particular, and more generally
in the Middle East and the eastern Mediterranean Basin.

There have been a number of books published over the past decade on the
topic in specific countries, including Mediterranean and arid climate regions,
such as Australia (Parson & Cuthbertson 2001, Muyt 2001, Hussey et al. 2007,
Richardson et al. 2011), South Africa (Henderson 2001) and California (Bossard
et al. 2000). This book is not unique, but it is the first that provides a general
overview of the major invasive alien plants that can be found today in natural
areas in Israel.

In view of the rapid dynamics of biological invasion, this edition will probably
have to be updated in the near future.
 





 



 



 


   
  

 

 
   
    
   
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1 – Invasive Plant Species: Basic Concepts and
General Information

1.1 – Alien species, invasive species and invasion stages

Pyšek et al. (2004) defined “alien species” as “species, subspecies or lower
taxa introduced outside of their natural range (past or present) and outside of
their natural dispersal potential. Their presence in the given region is due to
intentional or unintentional introduction or care by humans, or they have arrived
there without the help of people from an area in which they are alien”. According
to this definition, in Israel, any species that does not belong to the native flora,
or in other words, to the complex of plant species growing in our region without
human intervention (see Danin 2004 and Fragman et al. 1999) is considered an
alien species.
Most of the alien species in Israel are cultivated plants, sometimes called “exotic
species”, that is, species introduced deliberately to Israel from other parts of the
world, usually for landscaping purposes. A detailed list of alien cultivated plants
15
introduced to Israel can be found in the “Guide to cultivated plants in Israel”
(Fahn et al. 1999) and in the list published by the Israel Ministry of Agriculture
(Galon et al. 1993). The Ministry of Agriculture is now preparing a new list of
ornamental plants in Israel together with the Jerusalem Botanical Gardens.
The definition of a plant as “alien” is independent of its ability to reproduce
without human intervention in the region to which it was introduced and is not
related to potential damages it can cause to local ecosystems. These criteria
are considered only when defining a plant, or any other organism, as invasive.
Thus, according to Wilson (2003) an invasive species is a “species of plant,
animal, or microorganism that is both alien to the environment in which it lives
and destructive in some manner to the environment and its inhabitants” (Wilson
2003). It is very important to stress that “only a fraction of introduced species
successfully establishes or invades in the new region” (Pyšek & Richardson
2010). Therefore it is necessary to keep in mind that most alien species do not
become invasive and thus cause no harm to surrounding ecosystems.
The proliferation of an alien plant species is a process that occurs spatially
over a very variable period of time. Although some taxa can proliferate very
quickly in a new region, most invasive plants become established progressively
and it is therefore necessary to distinguish various proliferation stages in order
to determine the exact character and extent of invasion of an alien species.
Richardson et al. (2000, 2011) suggested differentiating between three main
invasion stages: casual, naturalized and invasive (table 1).
In addition to these three invasion stages another term has been proposed:
invasive transformers. This term describes the sub-group of invasive species
having the ability to “modify the traits, the conditions, the form or the character
of an ecosystem over a substantial area relative to the extent of that ecosystem”
(Richardson et al. 2000). “Transformer plants” cause changes in one or more
 Table 1. The invasion stages of alien plant species according to
Richardson et al. 2000

Invasion Stage Description

Invation Stage Description

Casual Alien plants that flower and occasionally reproduce, but cannot
create a sustainable population. Consequently the population
cannot survive in the absence of repeated human introduction of
new individuals.

Naturalized Alien plants that establish populations that renew themselves

without human intervention, for a consecutive period of at least
ten years.

Invasive Alien plants that are naturalized but produce a very large amount
of seed-bearing offspring that are dispersed over great distances
from parent plants. These plants have an invasion rate of more
16 than 100 meters from their parent plants over a period of 50
years, or at least six meters within three years in plants with
rhizomes.

Figure 1. Hottentot fig (Carpobrotus edulis), an invasive plant from South Africa that creates a
dense, thick layer of vegetation that reduces soil pH. The plant matter also modifies the level of
nitrogen and calcium in the soils. C. edulis is an “invasive transformer” in the coastal sands along
the Mediterranean shore of Israel.
Figure 2. Camphorweed (Heterotheca subaxillaris), an alien species from the United States
that began to spread in Israel’s coastal sands in the late 1980s. This species is an invasive
transformer that forms dense stands and stabilizes moving sands, thus completely transforming 17
the natural character of the habitat. Specialized native psammophilic plant and animal species
that require the bare landscapes formed by sand dunes are displaced by camphorweed.

of the physical components of the ecosystem they have invaded. These species
can, for example, modify soil properties by enriching it with nitrogen or reducing
pH (figure 1). Phenomena such as sand stabilization (figure 2) or increased
wildfire frequency1 are created by invasive transformer plants. In aquatic
ecosystems, e.g. rivers, marshes and lakes, the effect of invasive transformer
plants is usually extreme, as the coverage of water bodies by floating invasive
plants modifies the water temperature, reduces pH and leads to a serious drop
in the percentage of oxygen in the water (figure 3). Most invasive aquatic plants
are transformer plants.

Figure 3. Water hyacinth

(Eichhornia crassipes) is
a floating aquatic plant
originating in South
America. It invades
bodies of standing or
slow-flowing water and
may completely cover
the water surface, as can
be seen in this photo,
taken in the western
Sharon plain (north of
Tel-Aviv), modifying water
temperature, oxygen
concentration and pH.
Most invasive aquatic
plants are defined as
transformer plants.
1
Occurs in California, but has not yet been described from Israel.
 Most alien plant species in a given area will not become invasive. Despite the
great variability that exists between different areas, roughly 10% of the alien
plant species introduced to a given area become “casuals”; 10% of these
become “naturalized” and about 10% of these become “invasive” (i.e. 1% of the
“casual” species, or 1‰ of the ”introduced” species). These proportions, which
are the basis for the “tens rule” (Williamson 1993, Williamson & Fitter 1996)
should be regarded only as a general approximation. It expresses the belief that
only a small proportion of the alien species introduced to a given area could
become invasive (Richardson & Pyšek 2006). Moreover, according to Rejmanek
et al. (2005) about 10% of the invasive species become “transformer plants”.
However, according to the early results of a survey now being conducted by Ori
Fragman-Sapir and the author, the tens rule is not consistent with the situation
in Israel. Between 4000-5000 alien species have been introduced to the country,
but only 170 species have been defined as casual, naturalized or invasive, that
is to say 4% at the most. Nevertheless 34 of the 170, equaling 20% have been
defined as invasive species.

18
1.2 – A glimpse at the enemy release hypothesis
Although this book aims to focus on practical and applied aspects of invasive
plants in a specific region, namely Israel, it is necessary to mention briefly the
most basic hypothesis that account for the proliferation of an alien plant species
once it has been introduced to a new region, outside its natural range: The
enemy release hypothesis (ERH).

The enemy release hypothesis is probably the most well-known theory

mentioned in invasion ecology when addressing the issue of the successful
establishment of alien taxa in new habitats (Keane & Crawley 2002, Inderjit et
al. 2005). The ERH stipulates that an alien species, for example an alien plant,
when introduced in a new ecosystem, lacks natural enemies, e.g. herbivores or
pathogens, since these are generally not introduced with the alien plant. Since
the alien plant is not susceptible to local seed-eaters, such as insects, nor is it
vulnerable to local fungi, no organism can keep the alien individuals in check
and the alien taxa can proliferate. In other words the alien plant experiences
far fewer limiting constraints compared to native plant species that are under
permanent stress from local organisms.

The ERH is based on the fact that invasive plants are introduced to completely
different ecosystems than those they originally belonged to. The interactions
that exist between alien species and organisms in their original ecosystems
are lacking, due to the different species composition of the invaded ecosystem.
Thus, for example, the most famous alien invasive species in Israel, the golden
wreath wattle (Acacia saligna), grows wild only in southwest Australia. The life
expectancy of this species in its natural range is only 5-12 years, and rarely do
trees reach the age of 20. In Chile and South Africa, on the other hand, where
the species is exotic and invasive, wattles can live 30-40 years (O’Sullivan et al.
 2008). This difference results from the fact that in its natural range, in southwest
Australia, Acacia saligna is part of a local ecosystem and is affected by native
organisms that naturally regulate the tree population. A local rust fungus
species (Uromycladium tepperianum) causes the tree’s leaves to shed and
the tree to die (figure 4). A native beetle, the acacia seed weevil (Melanterius
compactus), which feeds on the tree’s seeds while they are still in their pods
(figure 5), limits the acacia’s reproduction. These two species regulate golden
wattle populations. When they are absent from the ecosystem to which the tree
was introduced, such as in the Mediterranean region in Israel, the wattle can
form large unregulated populations, which invade natural areas.

The ERH remains central in invasion ecology. Yet the absence of natural
enemies is not the only condition required for an alien plant to become invasive
(Thompson & Davis, 2011). In order to become successfully established in a new
ecosystem and then to proliferate, the alien plant taxon needs to have a suite of
biological and ecological traits that will enable it to compete with native species
and to cope with abiotic conditions that are sometimes significantly different
from those found in their original distribution area. 19
®PhotoÑ Wayne O'Sullivan ≤∞∞∑©

Figure 4. The rust fungus Uromycladium tepperianum on a golden wreath wattle in southwest
Australia. In its natural range this fungus acts as a natural enemy that reduces the trees life span
(Photo: Wayne O’Sullivan 2007).
 Figure 5. The acacia seed weevil Melanterius compactus feeds on golden wreath wattle seeds.
(Photo: Fiona Impson 2007).

20 1.3 – Invasive plants: general biological and ecological

characteristics
Although all invasive plants do not necessarily share exactly the same
characteristics, several biological and ecological traits are redundant among
most invasive taxa. One can point out at least nine biological and/or ecological
traits that are common to most of the invasive plant species found in Israel, and
in other regions. All the plant species given as examples below are invasive in
Israel.

(1) Ecological plasticity: the ability to grow and develop in a wide variety
of habitats is characteristic of many invasive plants. Water lettuce (Pistia
stratiotes) and water hyacinth (Eichhornia
crassipes) for example survive and
proliferate in Israel’s Mediterranean
region, despite low winter temperatures,
that occasionally drop below zero (°C).
This proves that water lettuce and water
hyacinth have great ecological plasticity,
which allows them to succeed in ambient
temperatures completely different than
those found in their natural range
– tropical South America (figure 6).
When the first foci of these plants were
Figure 6. Water lettuce (Pistia stratiotes) found in Israel in the early 2000s it was
photographed in the Hula Valley in February initially thought that the below freezing
2009. Contrary to expectations, low winter
temperatures of winter would create
temperatures in Israel did not cause
the collapse of water lettuce populations unsuitable conditions and eventually
in Israel. prevent invasion. Monitoring of several
 populations showed the ecological
plasticity of these taxa had been
underestimated.

(2) Asexual (vegetative) reproduction:

the ability to reproduce by means other
than seeds, e.g. creeping rhizomes or
cuttings, is fairly common in invasive
plants. This trait allows them to create
populations within short periods of
time, independent of pollinators and
other seed dispersal vectors. This is
found in most invasive aquatic plants,
e.g. African payal (Salvinia molesta),
red water fern (Azolla filiculoides), Figure 7. Bermuda buttercup (Oxalis
and in a considerable proportion of pes-caprae), an invasive plant from South
Africa that reproduces in Israel, but only
invasive herbaceous species, e.g.
vegetatively.
kikuyu grass (Pennisetum clandesti- 21
num), weakleaf bur ragweed (Ambrosia confertiflora), silverleaf nightshade
(Solanum elaeagnifolium), umbrella sedge (Cyperus involucratus) and
Bermuda buttercup (Oxalis pes-caprae) (figure 7). Several woody invasive
plants, such as tree of heaven (Ailanthus altissima), largeleaf lantana
(Lantana camara) and golden wreath wattle (Acacia saligna) are also able to
reproduce vegetatively.
(3) Rapid regeneration following damage: most invasive plant species
in Israel, and especially woody species, have the ability to regenerate
very rapidly following serious disturbance such as fires or cutting
(figures 8 and 9). This trait, combined with the other characteristics
mentioned above and henceforth, makes managing invasive plant
concentrations such a complex task.

Figure 8. Regeneration of golden wreath wattle (Acacia saligna, right) and willow wattle
(A. salicina, left), only a few weeks after cutting.
 Figure 9. Weakleaf bur ragweed (Ambrosia confertiflora) on the Alexander river bank
in the Sharon region, about two months after mowing.

22 (4) Rapid growth: most invasive plants are characterized by rapid growth.
This is particularly true for woody invasive plants in relation to native
woody species. For example, the tree of heaven can grow at a rate of 1-1.5
meters during one growth season (figure 10); the castor-oil plant (Ricinus
communis) can grow at a rate of 2 meters annually. Virtually no native trees
and shrubs have comparable growth rates in Israel.
(5) Competition: invasive plants use local natural resources, such as water
in the soil, very efficiently, and can thus successfully compete with native

Figure 10. Tree of heaven (Ailanthus altissima) understory. Thanks to its rapid growth, this
species, which originates in China, was chosen for landscaping in gardens and streets of many
cities in Israel. The blue arrow marks the annual growth. There is no native tree species in Israel
that has a similar growth rate.
 Figure 11. Invasive golden crownbeard (Verbesina encelioides) in the Kadima Nature Reserve in
the Sharon region. Its allelopathic traits provide it with a significant competitive advantage that
allows it to establish itself and invade natural areas where
23 it displaces native plants. 23

species. The mesquite (Prosopis juliflora) that invades arid areas in Israel
and in Jordan, develops a broad horizontal root system as well as deep roots
that reach depths of 15-20 meters. This allows the tree to exploit water from
a large volume of soil, at the expense of native species. The same applies to
the golden wreath wattle (Acacia saligna) which has taken over large areas
in the dry habitat formed by the coastal sand dunes. Many invasive plants
also have improved competitive abilities due to allelopathy, which allows
them to release chemical compounds
that prevent seed germination in native
species. This trait is found, for example,
in the tree of heaven (Ailanthus
altissima), the western coastal wattle
(Acacia cyclops), the common thorn
apple (Datura stramonium), the largeleaf
lantana (Lantana camara) or in the golden
crownbeard (Verbesina encelioides), an
invasive species that is found in Israel in
nature reserves in the Sharon region and
in the Agur Sands Nature Reserve in the
Negev (figure 11).

(6) Early seed production: the ability to

produce seeds early in its life cycle (a
short time after germination) is a major
trait among invasive woody species,
particularly trees. The golden wreath
wattle (figure 12) can produce seeds two
Figure 12. Ripe seeds of golden
wreath wattle (Acacia saligna) in their
pods a short time before dispersal.
This species can begin producing
seeds as early as the two years after
germination.
 years after germination (Richardson & Kluge
2008); the mesquite begins flowering and
seed production as early as the age of three
to five years (Pasiecznik et al. 2006). New
individuals thus become seed bearers within a
short period of time and rapidly enlarge the
soil seed bank in infested habitats.

(7) Various means of seed dispersal:

Invasive plant species often have a number of
methods of dispersing seeds, which ensures
rapid and efficient distribution. In many
cases their seeds are dispersed by wind and
by water – runoff or water flowing in streams.
Figure 13. A local ant species
In addition, seed dispersal in some invasive
(Messor semirufus) carrying a seed species occurs via local invertebrates such
of Verbesina enceloides in the Negev as ants (figure 13), or vertebrates such
desert.
24 as birds, bats or other mammals, which
disperse seeds over large distance. Fruit
bats, for example disperse the seeds of the chinaberry (Melia azedarach) in
the Mediterranean region of Israel. Tristram’s grackle, a native Israeli bird,
disperses seeds of three invasive Ficus species in the area of the Ein Gedi
Nature Reserve in the Dead Sea valley (figure 14). Fruits of species such

Figure 14. Tristram’s grackle, a native Israeli bird that lives in the Dead Sea valley, disperses
seeds of three invasive Ficus species in Israel, including the Indian banyan (Ficus benghalensis
pictured here). Dispersal occurs in the Ein Gedi Nature Reserve and along nearby wadis. Seed
viability is usually not affected by passage through the bird’s digestive tract. On the contrary, this
process accelerates seed germination and they germinate immediately after they are dropped.
 as weakleaf bur ragweed, rough
cocklebur (Xanthium strumarium,
figure 15) and spiny cocklebur
(Xanthium spinosum) cling to the
fur of native mammals such as
jackals and foxes, and are thus
dispersed over long distances.

(8) Producing large quantities of

seeds: many invasive species
produce extremely large quantities
of seeds. This trait combined with
the lack of significant predation on
the seeds, causes large quantities
of viable seeds to accumulate in Figure 15. Rough cocklebur (Xanthium
strumarium), an invasive species from
the area the plant invades. In
South America, produces numerous burrs
this manner most invasive plants with hooked spines that cling to the fur
create a much larger “seed banks” of mammals thus enabling long-distance 25
than do native species. Mature dispersal. The burrs also float on water and
are therefore readily transported over long
individuals of taxa such as tree distances.
of heaven (Ailanthus altissima) or
tree tobacco (Nicotiana glauca) can produce hundreds of thousands of viable
seeds each season. The invasive Australian acacias, such as the bramble
wattle (Acacia victoriae) and the western coastal wattle (Acacia cyclops)
(figure 16), produce seed banks with thousands of seeds per square meter.

Figure 16. Most of the invasive Australian acacias produce large numbers of seeds. On the right
are pods of bramble wattle (Acacia victoriae) that recently invaded the northern Negev and the
southern Mediterranean region in Israel. On the left are open pods of the western coastal wattle
(Acacia cyclops) that at this time is invasive principally in the Judean Mountains. The number of
seeds found in the soil beneath mature western coastal wattle trees can reach up to 5900 seeds
per square meter.
 (9) Extended seed dormancy: this trait is common to a considerable number
of invasive plant species and characteristic particularly of legume seeds
such the various Acacia and Prosopis species. Some of the seeds, instead
of germinating immediately after dispersal, remain in the ground, still
viable for periods ranging from a few to dozens of years. This trait allows
them to germinate long after dispersal. Golden wreath wattle (Acacia
saligna) and western coastal wattle (Acacia cyclops) seeds can preserve
viability for at least 50 years (Holmes 1989, Richardson & Kluge 2008). The
dormancy period of most invasive plants is still not known with certainty,
as this requires long-term research. Information relating to this property is
very important in order to develop efficient control strategies for infested
areas, especially when eradication is the goal, as removing all the existing
specimens in a given area is not sufficient to ensure eradication. As long as
dormant viable seeds remain in the ground they can germinate, even after
long periods of time and thus renew the invasive population.

26 1.4 – The dynamics of invasive plants: Basic aspects and

examples from Israel’s alien taxa
In the past decade several studies have addressed the issue of invasive plants
dynamics over space and time (see ref. in this section). Existing information
relating to the topic is crucial, as it enables us to evaluate what might be the
trend of alien plant proliferation within a given region in the near future. Two
main lessons can be drawn from the studies that focused on invasive plant
dynamics:

Firstly, a number of studies indicated that the probability of invasion increases

with the length of time that has elapsed since the alien species was introduced
to a new area (Richardson & Pyšek 2006, Rejmanek et al. 2005). Thus, if an
alien plant species is found in a given area for a long time and has therefore
become acclimated to local conditions, the probability of it becoming invasive
is higher than if it is in the area for a short period of time (Pyšek & Jarosík
2005). According to this statement, which seems to be widely accepted
among invasion ecology specialists, one should expect an increase in the
number of invasive plants in Israel in the coming decades since the bulk of alien
plant taxa were relatively recently introduced to the country. Most alien plant
species were introduced to Israel only during the 20th century, i.e. later than
in countries such as South Africa or Australia where invasive plant taxa are
numerous.

Secondly, the invasion rate of an alien plant species is not linear. The dynamics
of the invasion process of alien species in a given area is characterized by two
successive stages: during the first stage or lag phase the rate of spread is
slow and consequently there are few new individuals or foci. Alien plant
species at this stage are generally regarded as “naturalized” rather than
“invasive” (see table 1). In the second stage or exponential phase, the alien
species accelerates its rate of invasion and consequently begins to form
large populations and numerous concentrations throughout the given area
(Whitney & Gabler 2000, Richardson 2001, Groves 2006, and Richardson
& Thuiller 2007). Plant species that have reached this stage are considered
“invasive” (see table 1).
The lag phase is also called the minimum residence time. The length of this
period is not constant and varies significantly between species. European studies
have shown that the minimum residence time, in which the rate of invasion is
slow, can last for several dozens of years in shrub species, and an average of
170 years or more in tree species (Richardson 2001, Pyšek & Jarosík 2005). In
Germany there were reports of alien tree species that became invasive after
more than 300 years from the time they were introduced to the country (Groves
2006). The transition from the lag phase to the exponential phase is sharp and
fast. The factors underlying this sudden change, i.e. the transition from the
first to the second stage are not completely understood. Richardson (2001)
identified four potential factors that can explain the transition:
(1) Genetic adaptation of individuals in the alien species population that provide 27
them with improved resistance to local conditions, the ability to survive and
perform optimally, and to produce large amounts of viable seeds.
(2) Arrival of an alien organism to the invasion area that enables pollination or
dispersal of an alien plant species already established in the area. This can
explain the invasion dynamics of a number of Ficus species that are pollinated
only by specific pollinator wasp species (Richardson et al. 2000b).
(3) The sharp increase of disturbed areas in the last decades as a result
of anthropogenic development, has indubitably created an unprecedented
multitude of opportunities for invasive plants to proliferate since most
invasive plant species first establish themselves in disturbed habitats.
Therefore the number and the extent of infested areas grow accordingly.
(4) Consequently, the rate of seed production and seed bank accumulation in
infested areas grows exponentially.
(5) In addition to these, there is another possible explanation: Changes in the
behavior pattern of local animals that begin to consume and disperse the
seeds of alien species. This change, that is difficult to predict, could partially
explain the transition from the lag to the exponential phase especially when
seed dispersal by native bird species is involved.
Figure 17 presents five examples of invasive plants in Israel that invaded the
country at different times and entered the exponential phase during the past
years. The golden wreath wattle (Acacia saligna) and the western coastal wattle
(Acacia cyclops) were introduced to Israel in the early 1920s by the British, for
afforestation and soil stabilization. The golden wreath wattle became invasive
only towards the late 1970s, i.e. after 50 years. The western coastal wattle
became invasive only in the late 1990s – 75 years after it was introduced
to Israel.


 












 


  

















































28







 



  






Figure 17. Transition from the lag phase to the exponential phase in five invasive plant


species in Israel. The date of introduction to Israel (known or estimated) is marked


by triangles. The duration of the lag phase is marked with low density points whereas


the exponential phase is marked with high density points. The length of the lag phase
differs significantly between species and ranges between 15 years (camphorweed)
and 75 years (western coastal wattle).

The two other trees, the Indian banyan

(Ficus benghalensis) and the bramble
wattle (Acacia victoriae), were introduced
in Israel at least 50 years ago (the exact
date is unknown) and became invasive
only in recent years: the Indian banyan,
that invades only oases in the Dead
Sea Valley, became invasive in the
mid-1990s, and the bramble wattle
began to proliferate only from the end
of 2005 (figure 18). In these cases the
minimum residence time lasted 35-45
years. However some species may reach
the exponential phase within few years
following introduction: For example the
camphorweed (Heterotheca subaxillaris),
which was intentionally introduced to Figure 18. Invasion of bramble wattle
(Acacia victoriae) north of Be’er Sheva,
1975 in Israel, became invasive as early
on the fringes of the area where the tree
as 1990, that is only 15 years after its was once planted in the late 1980s and
introduction. early 1990s.
The fact that the invasion rate of alien plants is not constant but composed
of two stages – a lag phase, followed by an exponential phase – leads to the
almost definite conclusion that some of the “naturalized” species will become
“invasive” in the near future, i.e. they will reach the exponential phase.
Consequently, the number of invasive plants in a given area is expected to rise,
even if no additional alien plant species are introduced (Richardson & Pyšek
2006). According to Richardson (2011) the alien species which are still in
their lag phase, represent a “major invasion debt for decades and centuries to
come”.
Therefore, our current understanding of the dynamics of invasive plants does
not allow us to assume that the phenomenon of invasive plants will weaken and
wane in the near future.

1.5 – Examples of lists of invasive plants from

Mediterranean and arid regions in the world
Since the early 1990s various countries throughout the world have compiled 29
lists of invasive plant species that endanger native ecosystems. Below are
several examples of lists and databases created by national and international
authorities with the help of leading scientists in the field of invasive plants. Since
this book focuses on invasive plants in Israel most of the following examples
are from countries with Mediterranean or arid climatic regions, similar to those
found in Israel.

Invasive plants in Europe: EPPO and DAISIE

The European Union published a survey of invasive species that covers 49
European countries, including Turkey and Mediterranean Europe countries, as
part of the DAISIE (Delivering Alien Invasive Species Inventories for Europe)
project. The survey found 5789 alien plant species, of which 2843 were alien
to Europe (i.e. with a geographic origin outside the Old World). One of the
advantages of this database is that the current invasive stage of each species
is noted according to the terminology presented in table 1 at the beginning of
this chapter. The database also assesses the residence time of each plant on the
continent which is very valuable information (Pyšek et al. 2008).

Another database was compiled by EPPO (European and Mediterranean

Plant Protection Organization), an organization that publishes monthly
reports on observations of alien plants (and also of other alien organisms)
in European countries, the Maghreb (Algeria, Tunisia and Morocco), Jordan,
Israel and some of the central Asian countries – Azerbaijan, Uzbekistan and
Kirghizstan.

The California Invasive Plant Inventory

Cal-IPC, the California Invasive Plant Council was established in 1992. In 1994
the first list of invasive plant species in the state was compiled. This list was
 updated a number of times and in February 2006 a final report was published
(Bossard et al. 2006). The plant species were rated according to their ecological
threat level: high, moderate and limited. In addition, species with a high
invasive potential that are liable to invade new native habitats, trigger an alert
designation, which can be used for management purposes.

The 2006 report lists 193 species of alien plants that invade natural areas in
California. Of these, 39 have been rated high, 65 moderate and 89 still have
a limited threat potential at this time. Alert designations were triggered for
31 species. The next report, published in February 2007, already had seven
additional invasive species, two with a limited threat level, five moderate and
one high.

Invasive plants in Australia: Noxious Weeds, WONS, National

Environmental Alert List, Sleeper Weeds

As many as 27,000 species of alien plant species have been introduced to

Australia, of which 2779 are naturalized (Groves et al. 2005). Several lists of
30 invasive plant species according to specific needs and based on different criteria
have been compiled in Australia. The major lists are:

(1) A list of noxious weeds has been compiled for each of the Australian states
and territories. Originally the term “noxious” was used to indicate alien
plant species that caused damage to agriculture. Today, however, the term
is used to designate alien plants that invade natural areas and require
control programs to reduce their negative effect (Groves et al. 2003). In
2005, the list of noxious weeds included 429 species for all Australian states
and territories.
(2) The list of Weeds of National Significance (WONS) was published in 1999.
It includes 21 species that are all considered to pose a major threat to
Australian ecosystems. Among these species four are invasive in Israel, for

Figure 19. Jerusalem thorn

(Parkinsonia aculeata) in the
Bteha Nature Reserve north of
Lake Kinneret (Sea of Galilee).
 Figure 20. The karroothorn (Acacia
karroo), an acacia species native
to southern Africa, is already
considered a severe threat to
native ecosystems in Australia if
not immediately managed. The
increasing number of individuals
found just recently in natural sites
in Israel seems to confirm this
assessment.

example the Jerusalem thorn (Parkinsonia aculeata, figure 19) which, in

spite of what its vernacular name may suggest, is an alien taxa in Israel,
31
imported from Mexico.
(3) The list of sleeper weeds, published in 1999, lists alien weeds that for many
years seemed harmless, but in the wake of natural disasters such as wildfires
or drought were found to spread rapidly. The list includes 17 species, none of
which is known to be a casual, naturalized or invasive species in Israel.

(4) In 2000 the Australian Department of the Environment compiled a list of

alien plant species that although in early stages of invasion, i.e. in their “lag
phase”, had a high invasive potential, thus posing a severe threat to native
ecosystems if not immediately managed. This list, the National Environmental
Alert, includes 28 species, of which one species, the karroothorn (Acacia
karroo), is naturalized in Israel, (figure 20).

Invasive Plant Species in South Africa: SAPIA, Declared Weeds

and Invaders

In 1983 the South African government passed the Conservation of Agricultural

Resources Act, listing 198 alien plant species declared “weeds and invaders”.
The plants on the list were ranked in three categories with specific regulations
relating to commerce, planting and control:

a. Declared Weeds – category 1: these species are prohibited on any land or

water surface in South Africa. Plants in existing sites must be eradicated or
controlled. At least 122 species were listed in this category in all or part of
South Africa (Henderson 2001). Among these 12 are invasive in Israel.

b. Declared Invader – category 2: these species may not be imported or traded,

except by permit holders. The species in this category may be planted in
demarcated areas under controlled conditions. Any specimens found outside
these sites must be controlled and eradicated where possible. Planting is
 expressly forbidden along rivers or
in wetlands: “Declared invaders”
are prohibited within 30 meters
of the 1:50 year floodline. This
category includes 38 plants in
South Africa such as golden wreath
wattle, western coastal wattle and
the Murray red gum (Eucalyptus
camaldulensis), all invasive in
Israel.
c. Declared Invader – category 3:
species in this category may not
be planted or traded, but there
is no need to remove existing
populations, as long as measures
are taken to prevent further
spread. There are now 42 species
32
in this category in South Africa,
including the tree of heaven and Figure 21. Brazilian pepper-tree (Schinus
terebinthifolius) one of three invasive plant
the Brazilian pepper tree (Schinus
species included in the IUCN’s 100 of the
terebinthifolius, figure 21), both World’s Worst Invasive Alien Species list,
invasive also in Israel. which is found in Israel as well.

Between 1994 and 1998 the Southern African Plant Invaders Atlas (SAPIA) was
compiled. As part of this project 497 alien species were identified as “naturalized”
or “invasive” according to Richardson et al. (2000) and Pyšek et al. (2004, see
table 1). This survey is very interesting from an ecological viewpoint because
it focused on natural and semi-natural areas. The number of naturalized or
invasive species may be even greater, since the survey focused mainly on woody
species (Richardson et al. 2005). The number of alien herbaceous species in
natural areas in South Africa is probably higher than what appears in the SAPIA
survey.

The lists of invasive species mentioned above testify to the fact that invasive
species are an ecological concern of major importance in Mediterranean climate
regions and in semiarid areas (Pyšek & Richardson 2010).

The proliferation of invasive plants in natural and semi-natural areas is

a relatively recent phenomenon in Israel, especially when compared to other
Mediterranean-type and arid regions in the world (South Africa, Australia,
California). Prior studies addressed the issue of alien plants in Israel (e.g. Dafni
& Heller, 1990), although mainly from the perspective of agriculture weeds. The
information presented in the next chapter shows that alien invasive plants do
not spare local ecosystems and the most recent observations in the field do not
allow us to assume that the rate of invasion in Israel will slow down in the near
future.
2 – Invasive Plants in Natural Areas in Israel:
A Review

2.1 – The factors underlying the invasion of alien

plant in natural areas in Israel
The factors underlying the invasion of alien plant species in natural areas in
Israel can be divided into two main categories: Firstly, human activities that
resulted in the introduction of alien plant species into Israel (section 2.1.1
below). Secondly, the recent and intensive land use developments that created
opportunities for alien plants to establish themselves and eventually invade
natural areas (section 2.1.2).

2.1.1 – Anthropogenic activities that introduced invasive plant

species in Israel
Afforestation 33
Afforestation is one of the sources of introduction of invasive tree species
into Israel. This situation is common throughout the world (Richardson 1998,
Richardson & Rejmánek 2011), particularly in countries with Mediterranean or
arid climates (Kull et al. 2011, Le Roux et al. 2011).

Several facts should be mentioned when evaluating the role afforestation played
in the current situation of invasive trees in Israel:

Afforestation activity in Israel was initiated during the British mandate

(1917-1948). Tree planting was planned and methodically promoted by the
British administration in Israel, as it was in other areas under British control such
as Cyprus. Extensive information can be found in the reports produced during
that period (Sawer 1922, 1931). The forestry department of the Jewish National
Fund (JNF) also planted many forests throughout the country, both before and
after the modern State of Israel was established. The forestry department
continues its planting activities in the country up to the present, although the
extent of newly afforested areas has decreased in recent years.

When considering the role of forestry in the subsequent invasion of alien trees
in Israel, and obviously in other regions, it is important to keep in mind that
the phenomenon of biological invasions was almost unknown until the late
1950s, when Elton’s book was published in 1958. Only 20 years later, in the
late 1970s did awareness of the problem of invasive plants begin to emerge
among ecologists. Understanding the nature of the problem and the magnitude
of its consequences for ecosystems was only fully assimilated in the late 1980s
(Di Castri et al. 1989). Thus, in Israel and in other countries, the use of alien
tree species for afforestation, which eventually became invasive trees, was not a
result of negligence or lack of professionalism, but rather due to ignorance of the
existence of the phenomenon of invasive plants. Acacia saligna, for example,
 was still a recommended planting
species in Israel in the late 1970s
(Zafrir 1976).

Choosing to use alien tree species

for afforestation in a small country
where there are 79 native tree
species, adapted to a wide variety
of habitats, could seem odd. Yet, in
Israel as in other countries, alien
trees were preferred over native ones
because their growth rate is faster,
and therefore they could give
good results within short periods of
time.

Several alien tree species used for

afforestation or soil stabilization
34 in Israel were afterwards found to
have become invasive. For example
the Murray red gum (Eucalyptus Figure 22. A young sapling of Eucalyptus
camaldulensis) was planted in Israel camaldulensis in the Apple-ring Acacia Nature
for years in the Mediterranean region Reserve near Ashdod.

of the country and was not considered

an invasive species. Eventually field
observations showed that Eucalyptus
camaldulensis invades wetlands such
as river banks (e.g. the Nahal Tavor
and Tel Dan nature reserves) and
sandy habitats with a high groundwater
table (e.g. the apple-ring acacia Nature
Reserve near Ashdod, figure 22, and
the Kadima National Park.)
The Calabrian pine (Pinus brutia)
was also planted in extensive areas
in Israel’s Mediterranean region
and during the past ten years many
invasive Pinus brutia trees have been
found in the Tabor oak (Quercus
ithaburensis) park forest in the Alona
Nature Reserve located in the Ramat
Menashe region, east of Caesaria
(figure 23). The Tabor oak park forest is
a unique vegetation unit, characteristic
to Israel and northwestern Jordan, Figure 23. Invasion of Pinus brutia in a Tabor
in which pine species are normally oak wood park in Ramot Menashe.
absent, even from the early succession stages leading to its establishment. This
vegetation unit is typically made of a scattered tree layer and a species rich
herbaceous layer. Establishment of Pinus brutia in this type of vegetation unit
overshadows and eventually displaces the herbaceous species, thus altering the
structure and the composition of the Tabor oak park forest.

In the semi-arid northern Negev and in the arid region to the south, afforestation
was conducted for the purpose of soil stabilization, microcatchment creation,
preventing illegal building, “landscape improvement” and recreation. The trees
planted were species known for their resistance to dry conditions. Some of these
became invasive species, among them Australian wattles such as Acacia saligna
(that was also planted in sandy areas), A. salicina and A. victoriae, alien
Prosopis species such as P. juliflora and other species such as Parkinsonia
aculeata.

Nonetheless it should be noted that at this point only a relatively small

proportion of the alien tree species introduced into Israel for planting have
become invasive. Alien tree species such as Pinus pinea, P. canariensis,
35
Eucalyptus gomphocephala, E. occidentalis. E. hemiphloia or E. torquata were
planted over large areas in Israel (Reves 2008) and have not posed any threat
to local ecosystems so far.

Ornamental Plants

Ornamentals are a major source of invasive plants in general and particularly

in Mediterranean-type countries (Marco et al. 2010): Many alien invasive
plants In South Africa were imported and planted as ornamentals (Alston
& Richardson 2006). In Australia as much as 94% of the alien plants introduced
were brought in as ornamentals, and they now represent 66% of the naturalized
or invasive plant taxa in the country (Groves et al. 2005, Caley et al. 2007).
So it is not surprising that we find a significant proportion of ornamentals
among the naturalized or invasive alien plants in Israel. Of the fifty species
portrayed in this book, however, about twenty were introduced into Israel
as ornamentals for gardens and parks. These include Ficus benghalensis,
Ficus microcarpa, Ficus religiosa, Schinus terebinthifolius, Washingtonia
robusta, Ailanthus altissima, Lantana camara, Carpobrutus edulis and Cyperus
involucratus.

Although ornamentals are planted mainly in urban areas, e.g. streets, parks and
gardens, some of them manage to “jump the fence” and to invade natural areas.
In Israel this process occur in two main ways:

The first one involves natural dispersal agents such as several native animal
species such as bats and birds feed on the fruits of ornamental plants and
disperse their seeds beyond the limits of the urban area. For example there are
observations from the past 20 years that the California fan palm (Washingtonia
robusta), and three alien fig trees (Ficus religiosa, F. microcarpa and
F. benghalensis) have been dispersed in the Ein Gedi Nature Reserve by local
 bird species (Blecher 2005, figures 24
and 25). Other ornamentals benefited
from additional dispersal agents
such as wind or runoff: For instance
the tree–of–heaven (Ailanthus
altissima) which was planted along
many streets in Jerusalem has
escaped the urban area and now
invades the Nahal Halilim Nature
Reserve near Jerusalem (figure 26).
The second way ornamentals invade in
natural areas is through garden refuse,
which is another common source for the
proliferation of ornamentals in natural
areas, particularly those reproducing
vegetatively. Throwing out stems
or rhizomes of alien ornamentals in
36 natural sites adjacent to residential
Figure 24. Individuals of Washingtonia
areas may be sufficient to produce robusta in the Ein Gedi Nature Reserve.
new infested sites. For example
numerous foci of the Hottentot fig
(Carpobrotus edulis) were created
this way along Israel’s coastal plain in
sandy habitats. The same occurs more
and more frequently when aquarium
or fish pond contents are dumped in
water bodies such as streams, canals,
swamps and ponds. In this manner
ornamental water plants such as
Eichhornia crassipes, Pistia stratiotes,
Salvinia molesta, Azolla filiculoides
and Myriophyllum aquaticum rapidly
infest new sites throughout the
Mediterranean region of the country.
This proliferation pattern is common
to many regions in the world (Bossard
et al. 2000, Muyt 2001, Romanowski
2011). Natural sites with stagnant or
slow flowing waters are particularly
vulnerable to aquatic ornamentals.
Figure 25. A young sapling of Ficus
Since the early 2000s the number benghalensis near a waterfall in the Ein
of natural sites infested by floating Gedi Nature Reserve. During the past 15
invasive ornamentals is constantly on years hundreds of individuals of this species
germinated along the watercourses in the
the rise. Unfortunately most species
Ein Gedi area. Apparently winter floods
are still sold in nurseries with no cannot prevent the establishment of the alien
regulation (figure 27). fig tree.
 37
Figure 26. Invasion of Ailanthus altissima in the Nahal Halilim Nature Reserve west of Jerusalem.

Other sources of introduction

In addition to afforestation and
ornamental plants, there are other
specific sources that can explain the
invasion of some alien species in
Israel’s natural areas: For example
the pop saltbush (Atriplex holocarpa)
was imported from Australia in the
early 1960s as a potential pasture
plant for Israel’s arid region, along
with the bluegreen saltbush (Atriplex
nummularia) that also originates
in Australia. The attempt failed but
meanwhile the pop saltbush spread
throughout the northern Negev to the
Arava and the Dead Sea Valley. It is
interesting to notice that the bluegreen
saltbush did not become invasive.

Another example is the Chinese

waterspinach (Ipomoea aquatica),
which originates in southern China.
This morning glory species is a common
edible plant in Southeast Asia. Workers
from this area often grow the plant in
Figure 27. Specimens of Pistia stratiotes
and of Salvinia molesta for sale in a plant
nursery in central Israel. Both species are
known as invasive species in many parts of
the world and have already been found in
natural areas in Israel.
 the foreign countries they work
in, e.g. Florida, Texas and Israel.
Since this species reproduces
vegetatively and grows very fast
it proliferates readily in wetlands.
A small population of this species
was found in the Harod River in
the eastern lower Galilee in 2009
and in the northwestern Negev,
near Kibbutz Nir Oz, in late 2011
(figure 28).

The particular case of the

camphorweed (Heterotheca
subaxillaris) should be also
mentioned since it was
deliberately introduced in Israel
38 in 1975 in order to stabilize
moving sands. Within two Figure 28. The Chinese waterspinach (Ipomoea
decades the plant became one aquatica) in Harod River, eastern Lower Galilee, in
of the worst invasive species autumn 2009.

along the coastal sands along

the Mediterranean shore and is
one the few alien plants known
as ‘invasive’ only in Israel.

The manner in which invasive

plants reach Israel is not
known for all the species found
here. Some probably arrived
in seed mixes for agriculture
or as food for fish and birds.
The burr ragweed (Ambrosia
confertiflora), which appears to
be one of the most threatening
invasive plants in Israel, and
the crown beard (Verbesina
encelioides, figure 29) probably
arrived in this way before they
began spreading to natural
areas.

Figure 29. Crown beard (Verbesina encelioides) at

the edges of an agricultural field in the northwestern
Negev. The plant probably reached Israel in a seed
mix intended for agriculture. The species is now
invasive in natural areas, such as the Agur Sands in
the Western Negev.
2.1.2 – Factors that increased the invasibility of
Israel’ natural ecosystems in recent decades
The invasibility of a region reflects its vulnerability to invasion. Several factors
determine the invasibility of a region such as climate, properties of native
species, natural ecosystems resistance to invasion and degree of disturbance
resulting from human activities (Lonsdale, 1999). Since invasibility is strongly
affected by the intensity of anthropogenic disturbances, the overall susceptibility
of natural local ecosystems to alien plant invasion may change significantly and
rapidly over time.

During the past 30-40 years Israel became a densely populated, industrialized
country with modern agriculture. This can largely explain the growing number
of invasive plant species found in the remaining natural areas, as well as the
steady increase in the extent of infested areas during recent decades.

Population density
Israel is a small, densely populated country. Its area is 22,000 km2 and its
population 7.5 million (in 2010). Population density is about 340 inhabitants 39
per km2 on average, and over 500 per km2 in the center of the country. One of
the consequences of increasing population density is the expansion of built up
areas, inevitably at the expense of natural areas. The extent of natural areas
in Israel has shrunk considerably within the past 50 years and in the most
populated areas, such as the coastal plain, remaining natural sites, even if
protected are very small. Many nature reserves, principally in the coastal plain,
comprise only a few hectares and even less (“flower-pot reserves”). For example
the total area of the Kadima Reserve is less than one hectare (0.9 Ha) as is that
of Tel Yitzhak Reserve (0.8 Ha). Due to their very restricted size these remaining
natural sites, surrounded by densely populated built-up areas are now ecological
islets extremely sensitive to invasive plants, since even small foci with relatively
few individuals, can rapidly cover a considerable part of the remaining natural
area.

At the same time, an increase in population density implies more disturbances

and more alien plants, particularly ornamentals, among which are potential
invaders. For example a few dozen individuals of the largeleaf lantana shrub
(Lantana camara) established themselves at the edge of the Coastal Iris Nature
Reserve, which was intended as a refuge for both very rare and endemic species
such as the coastal iris (Iris atropurpurea) and the Tel-Aviv garlic (Allium tel-
avivense), and considerably reduced its already very small dimensions (4.85
Ha). The same is true for the Liman Nature Reserve, a small (5.2 Ha) reserve
located in the northwestern Galilee coastal plain on calcareous sandstone
(named ‘kurkar’). In this reserve there are several very rare plant species
such as the Lesser Adder’s tongue (Ophioglossum lusitanicum) and the coastal
endemic Beirut Mullein (Verbascum berytheum). The Lesser Adder’s tongue, a
small native pteridophyte (5-8 cm height), is easily overtopped by the spreading
patches of the invasive Bermuda buttercup (Oxalis pes-caprae).
 This situation is not exclusive to Israel; a similar phenomenon and its associated
negative effects were identified in coastal areas in Greece (Arianoutsou et
al. 2010) and described also in the Cape Province in South Africa (Alston &
Richardson 2006).

Infrastructure expansion and habitat fragmentation

During the last twenty years the expansion of infrastructures, and particularly
the extent and the density of roads and railways networks have significantly
increased in Israel. This phenomenon played a role in the proliferation of
invasive plants throughout the country according to a well-known process that
has also been observed in other parts of the world: roadsides become preferred
habitats of many invasive plant species as they are moist, rich in nitrogen
and disturbed (Milton & Dean 1998, Arevalo et al. 2005, Flory & Clay 2009).
Consequently Acacia saligna, Parkinsonia aculeata, Nicotiana glauca and Ricinus
communis are conspicuously present along roadsides and railway tracks in
Israel’s Mediterranean region (figure 29). In the Negev species such as Prosopis
40 juliflora, Nicotiana glauca and Acacia salicina have also become established on
roadsides.

This situation is not an ecological problem on its own, since roadsides are
habitats that have been destroyed and lost their ecological value even before
invasive plants became established on them. However, roadsides can serve as
potential corridors used by invasive plants to spread into natural areas (Christen
& Matlack 2006, 2009, Gelbard & Belnap 2003, Kalwij et al. 2008). Roadsides
also serve as a stable seed bank from which new foci of invasive plants can
become established in adjacent natural areas.

The increase in the density of transportation infrastructures worsens the

fragmentation of natural habitats which consequently become prone to the
spreading of invasive plants. This process, together with the increase in
population density, has probably significantly intensified the proliferation of alien
invasive plants in Israel in recent years.

The lack of buffer areas between agricultural and natural areas

In a very small and densely populated country such as Israel it is not surprising
to find that numerous protected natural areas are contiguous to areas of
intensive agriculture. The absence of buffer areas between nature reserves and
cultivated areas is another spatial feature that accelerates the proliferation of
invasive plant species within natural areas in Israel. This situation is a direct
outcome of the shortage of land in the country, which is expressed by the
lack of buffer areas around natural areas such as are generally found in larger
countries with lower population densities. In Israel many protected areas and
reserves are both small and surrounded by intensively cultivated land. As a
result, alien plant species that were once known as weeds and found only in
cultivated areas, now invade nature reserves in various parts of the country.
Figure 30. Invasion of Acacia saligna along highway no. 1 in the Judean Mountains.
The tree has now infested a large part of the Judean Hills National Park, which is crossed
by the highway.
41

These include the silverleaf nightshade (Solanum elaeagnifolium), the fleabane

(Conyza albida), the Canada horseweed (Conyza canadensis), the asthmaweed
(Conyza bonariensis), the American pokeweed (Phytolacca americana,
figure 31), the crown beard (Verbesina encelioides) and the burr ragweed
(Ambrosia confertiflora). These species that normally are limited to agriculture
lands (figure 29) are now found in the natural areas contiguous to the cultivated
fields. For example the crown beard (Verbesina encelioides), a typical agriculture
weed is now invading the Kadima Nature Reserve in the Sharon region
(figure 11) and the edges of the Agur sands Nature Reserve, in the northwestern
and western Negev desert .

Figure 31. Invasion of Phytolacca americana in the Hula Nature Reserve. This alien species is
characteristic of field edges, but is now spreading into the natural area of the Nature Reserve.
 2.2 – The major effects of invasive plants on natural
ecosystems in Israel
The invasion of alien plants has two major ecological effects on local ecosystems:
(1) displacement of native plant species and (2) modification of the abiotic
characteristics of the infested habitats with adverse consequences for the whole
native biota. At this point it is not possible to clearly identify changes in fire
regimes in Israel that would result from the proliferation of invasive plants, as
happens, for example, in California with the invasion of Bromus tectorum.

2.2.1 – Displacement of local plant species

Due to their biological and ecological characteristics (see section 1.3) invasive
plants can establish themselves in new habitats. When they succeed in forming
dense stands, they gain supremacy over native plant species. As a result, local
plant species are progressively displaced from their habitat by the invader. This
process can be achieved through different strategies.
42 The most common method of displacement is simply by reducing access to
light for the native species. Once established, invasive plants rapidly form
dense patches that overtop, crowd
out and eventually suppress native
species by shading them out.
Consequently, local species are
almost completely absent from the
understory of most invasive trees
and shrubs. Native species beneath
the canopy of dense thickets
formed by Australian wattles in
Israel are quite rare, whatever
the type of the infested habitat
(figure 32).

Another method by which invasive

Figure 32. Understory of Acacia saligna in
a sandy loam habitat in the coastal plain in
the region of Netanya. In its natural state, this
habitat is very rich in herbaceous species, some
endemic to the coastal plain. Yet in infested
areas the dense tree cover creates shade that
significantly reduces the diversity of native plant
species.
plants displace local plants is with
chemical compounds that prevent
germination of native species in the
vicinity of the invading species. This
trait, known as allelopathy, is very
common in invasive plant species.
A number of invasive plants in
Israel possess this character, such
as Verbesina encelioides, Ailanthus
altissima, Acacia cyclops, Datura
stramonium, Nicotiana glauca,
Phytolacca americana, Schinus
molle, Schinus terebinthifolius,
Prosopis juliflora, Solanum elaeagni-
folium and Lantana camara (figure
33).

Competition for soil resources,

particularly water, also displaces
local species (figure 34). With
the exception of sandy habitats,
competitive displacement of local
species as a result of competition
for soil nutrients is not significant,
as most soils in Israel develop from
nutrient-rich sedimentary rocks.

Displacement of local plants by

invasive species is a gradual process.
The damage is mostly indirect and
expressed by a slow but constant
reduction of the extent of the habitat
available for the native species. In
terms of biodiversity conservation
this process becomes even more
serious when alien species invade
habitats populated by endemic
plants whose area of occupancy is
naturally very restricted. In these
cases the displacement process can
43
Figure 33. Understory of Lantana camara in
the small coastal iris Nature Reserve in the
coastal plain. This invasive shrub displaces
local species by creating dense thickets and
releasing chemical compounds from its leaves
that prevent local plants from germinating.
The understory of Lantana camara is totally
devoid of native species.

Figure 34. A specimen of Heterotheca subaxillaris displaces a specimen of Artemisia

monosperma, a native species (marked by blue arrow) in sand dunes along Israel’s coast.
 lead to a drastic reduction in the number of individuals and eventually to the
collapse of the entire population within a short period of time. The invasion of
coastal habitats by the Hottentot fig (Carpobrotus edulis) is one such case: in
a number of spots the species is displacing individuals of the coastal iris (Iris
atropurpurea), a native rare (‘red’) species endemic to Israel (figure 35) and
considered critically endangered by the IUCN (Shmida & Polak 2007).

2.2.2 – Invasive transformers

Among invasive plants some are able to modify, sometimes radically, the
character of the ecosystem they invade, especially its abiotic features. These
invasive plants are termed “transformers” (Pyšek et al. 2004). Several of the
most important invasive plants in Israel belong to this category: For instance
the Hottentot fig (Carpobrotus edulis) which threatens coastal habitats reduces
the soil pH and the concentration of nitrogen and calcium in areas it invades.
The golden wreath wattle (Acacia saligna) produces large quantities of leaf
litter that raises the nitrogen level in the soil thus affecting its nutrient balance.
This change encourages the development of other invasive plants that take
44 advantage of the high soil nitrogen level (Tye & Drake 2011).

Both Acacia saligna in the coastal sands of the Mediterranean region and Acacia
salicina on wadi banks in the Negev desert, modify the character of the natural
ecosystems by creating dense tree stands in areas whose natural vegetation
cover is open, and composed of low shrubs and herbaceous plants, rather than
dense tree thickets. This change, both in the composition and in the structure
of the vegetation cover in infested areas, severely impacts local animal species.
For example small mammals and birds, such as the greater Egyptian gerbil
(Gerbillus pyramidum) and the Eurasian stone-curlew (Burhinus oedicnemus),
are adapted to semi-stabilized sand habitats with low open vegetation that
provide them with a wide field of vision. A transformation of their habitat by

Figure 35. The invasive Carpobrotus edulis displaces the native and very rare Iris atropurpurea in
a protected sandy habitat near Netanya. The native geophyte (Iris atropurpurea) cannot compete
with the invasive succulent perennial (Carpobrotus edulis) and retreats from infested areas.
invasive ligneous species whose
cover results in dune stabilization
can negatively affect psammophile
species (Anglister et al., 2005). A
similar situation is created by the
invasion of Heterotheca subaxillaris
on the coastal sandy habitat along
the shore, where it sometimes forms
a closed, dense soil-stabilizing cover
(figure 36).

However the ultimate transformers

among invasive plants are aquatic
invasives, particularly free-floating
plants such as the water lettuce Figure 36. The camphorweed (Heterotheca
(Pistia stratiotes), the water hyacinth subaxillaris) is gradually covering the sands along
the coastal plain and literally transforming the
(Eichhornia crassipes), the Parrot
physical characteristics of this specific habitat
Feather Watermilfoil (Myriophyllum originally composed of patches of bare sand. 45
aquaticum), the red water fern (Azolla
filiculoides) and the Africa payal (Salvinia molesta). All of these species have
been spreading at an alarming rate in the past few years through the remaining
natural aquatic habitats in Israel. When these species invade stationary or slow-
flowing water bodies they form a dense layer or a compact continuous mat on
the water surface. The almost completely impermeable cover formed by these
floating invasives (figures 3 and 37) alters the physical characteristics of the
water body by decreasing oxygen concentration, lowering pH, modifying water
temperatures, and by increasing the accumulation of organic compounds in the
water through decaying plants. If the invasion becomes permanent the drastic
changes described above affect most of the vertebrates inhabiting the infested

Figure 37. The Gdora River (Western Galilee) choked by Eichhornia crassipes in 2009. The stream
banks are densely covered by Acacia saligna.
 water body and lead to their extinction (see examples in Bossard et al. 2000,
Parsons & Cuthbertson 2001).

2.2.3 – The habitats most vulnerable to invasion by alien plants in

Israel
In the array of natural habitat types found in Israel, two are particularly
sensitive and prone to invasion by alien plants: the sandy habitats, mainly along
the coastal plain, and the wetlands. The sensitivity of these habitats results
from the combination of their unique abiotic characteristics with the fact they
are probably among the natural habitats most disturbed in Israel over the past
50 years.

Sandy habitats
Sandy habitats in Israel are found in the Mediterranean region, along the
coastline, and in several areas in the arid region such as the large erg found in
the northwestern Negev desert, named the “Agur Sands”, the “Samar Sands”
area in the southern Arava Valley, or the eastern part of the Ramon crater in the
46 central Negev.
Sandy habitats along Israel’s coastal plain have been severely damaged by
land reclamation over the past 50 years. Construction of residential areas,
development of transportation infrastructures and sand quarrying (now illegal)
have inevitably led to the destruction and the fragmentation of the coastal
dunes along the shoreline. Nonetheless, several areas have been preserved
since they were declared Nature Reserves or National Parks. Yet, almost all
the remaining fragments of sandy habitats along the coastline, including
protected sites, have been invaded by alien plant species over the past 30-40
years, and are today heavily infested. Of the four most common species in
sandy habitats along the shoreline – the golden wreath wattle (Acacia saligna),
the camphorweed (Heterotheca subaxillaris), the Hottentot fig (Carpobrotus
edulis), and the beach evening primrose (Oenothera drummondii) – the first
three are considered “transformer” species (see section 2.2.2, figure 38). The
camphorweed, the Hottentot fig and the beach evening primrose invade almost
exclusively sandy habitats in Israel while the golden wreath wattle can be found
in a variety of habitats in the Mediterranean region of the country. Moreover,
in terms of population sizes, a recent countrywide survey has shown that the
coastal sands are populated by the largest invasive plant populations in Israel
(Dufour-Dror 2009).
The sensitivity of coastal sand dunes to invasions is not a unique Israeli
phenomenon: studies have shown a similar situation in South Africa (Gaertner
et al. 2009) and in Portugal (Marchante et al. 2008) mainly resulting from
invasion by Australian acacias.
One explanation for the unique sensitivity of coastal sandy habitats to the
invasion and establishment of invasive plants lies in the abiotic character of this
habitat, which is determined mainly by the type of soil. Because of their soil
 47

Figure 38. The five most frequent invasive plant species found in sandy habitats along the
coastal plain. Clockwise from the upper left: Acacia saligna, Carpobrotus edulis, Oenothera
drummondii, Oxalis pes-caprae and Heterotheca subaxillaris. The first two and the fifth are
transformer species.
 texture that cannot retain humidity, coastal sand dunes generally form drier
habitats. This is especially true for the Mediterranean climate zone, with the
exception of areas where the water table is particularly high. Moreover, sandy
soils are acidic and poor in nutrients (i.e. in basic cations). Therefore this type
of habitat typically is characterized by pronounced abiotic constraints compared
to other adjacent natural terrestrial habitats. A number of shrubs and native
herbaceous species in particular, are adapted to these conditions, and when
undisturbed they create an open, discontinuous vegetation cover with patches of
bare sand. Alien plants that are able to adapt to these limiting abiotic conditions,
readily establish themselves in open sandy patches, which could be considered,
to some extent, as ecological niches not filled by the native plant species. The
naturally open, i.e. discontinuous, vegetation cover in coastal sandy habitats
could explain why a number of alien species have so successfully invaded this
type of habitat in Israel. This assumption remains yet to be proven, since the
degree of man-induced disturbances in the establishment of alien invasive
species in sandy habitats might be more significant than previously thought.
Nonetheless, the ecological consequences of the spread of invasive plants in
48
sandy habitats along the coastal plain are particularly serious for three reasons
mentioned in short below:

First and foremost, coastal sands along the Mediterranean shore of Israel form, in
their natural state, a restricted habitat, limited to a narrow strip, a few hundreds
of meters wide, along the coastline. In view of the spreading rate of the various
invasive plants that have proliferated during the past 30 years in this type of
habitat, we cannot rule out a scenario in which all the sandy habitats along the
coastal plain will be severely infested with invasive plants in the near future.
The second reason that makes the takeover of sandy habitats by invasive
plants particularly serious from an ecological viewpoint comes from the ability
of Acacia saligna, Carpobrotus edulis and
Heterotheca subaxillaris to completely
transform the basic character of the
coastal sand dunes (figures 39a, 39b and
39c), mainly by creating a continuous
vegetation cover where native plant
species are crowded out and eventually
displaced from large areas. This situation
is not common as yet in other natural
habitats in Israel, with the exception of
free floating invasive plants that may
completely cover water bodies.
Figure 40. A young specimen of The third reason is a consequence of the
Heterotheca subaxillaris (green rosette two previously mentioned: several rare or
in photo) that displaces Rumex occultans
(red flowers in photo), a species endemic
endemic plant species are found only in
to the coastal sands in Israel and the coastal sands (figure 40). Therefore
Lebanon. coastal sands constitute a unique

48
A

49

Figures 39 a, b, c. These
three photos were taken
in a sandy habitat south
of Netanya in the coastal
plain. The sites in each of
the pictures are only about
B 50 meters apart. Figure
38b shows a patch of bare
sand, semi-stabilized, with
scattered specimens of
Artemisia monosperma
and Retama raetam, both
native species; this photo
shows what the sandy
habitat looked like before
the coastal plain was
invaded, mostly by Acacia
saligna and Heterotheca
subaxillaris. Figure 38a
shows a population
of H. subaxillaris that
completely covers the soil
surface. Figure 38c shows
a dense stand of A. saligna
with specimens reaching a
height of 5 meters.
C
 habitat within the Mediterranean region, into which many desert species have
penetrated, some of which created unique ecotypes2. The presence of desert
species well beyond the limit of the arid region, in the area of the Mediterranean
basin is an example of azonal distribution (i.e. one that is not a consequence
of climatic conditions, but of soil conditions in this case) that increases local
biodiversity. Therefore, the proliferation of invasive plant species in this type of
habitat may pose a direct threat to several native species that are naturally rare
and endemic to this particular habitat.
A last aspect that should be mentioned about the vulnerability of sandy habitats
along the shore in Israel is the threat posed by another Australian acacia, the
western coastal wattle (Acacia cyclops), which is already invasive in Israel’s
mountainous area. This species has become one of the most problematic
invasive species in sandy habitats along the South African coast in the Cape
Province which has a Mediterranean climate very similar to the one in Israel
(Holmes et al. 1987, Impson et al. 2004, Kull et al. 2011). There is therefore, a
serious risk that this wattle species will begin spreading through the remaining
sandy habitats in Israel’s coastal plain in coming years. If this scenario were to
50
come true, the ecological damage to the sandy habitats along the Mediterranean
coast of Israel could be worse than any occurring up to now.

Sandy habitats in Israel have been severely affected by construction along the
coast. Direct destruction of these areas has been the major reason for the loss of
this unique habitat for dozens of years. It now seems that invasive plant species
have taken its place as the greatest threat to coastal sands, as they affect and
destroy the sandy ecosystem even in areas protected from development.

Freshwater and wetland habitats

Freshwater and wetland habitats in Israel include aquatic environments such

as streams, lakes and springs, whereas wetlands sensu lato comprise riparian
habitats such as stream or wadi banks, lake shores, as well as seasonal winter
pools and natural areas with a high water table. Freshwater ecosystems
and wetlands are well known for being particularly prone to invasive plants
(Richardson et al. 2007, Esler et al. 2008, Holmes et al. 2008, Catford et al.
2011) and regional surveys generally show a higher concentration of alien
invasive taxa in these habitats (Pyšek et al. 2010, Arianoutsou et al. 2010).

It is therefore not surprising to find a relatively large proportion of invasive

plant species in aquatic habitats and in wetlands in Israel. As seen in
table 2, at least 25 different species of alien invasive plants can be found in
these types of natural habitats in Israel. Of these, five species are free floating
plants considered “transformers”: Azolla filiculoides, Myriophyllum aquaticum,
Pistia stratiotes, Eichhornia crassipes and Salvinia molesta.

Like the sandy habitats along the coastal plain freshwater habitats and wetlands
occupy only small areas in Israel. Yet, large concentrations of invasive plants
2
For example, Retama raetam, Salvia lanigera and Moltkiopsis ciliata.
Table 2. A list of the major invasive plant species found in freshwater
and in wetlands in Israel (some of the species invade dry habitats
as well).

Freshwater
Species Wetlands
ecosystems

Azolla filiculoides ´ ≠
Eucalyptus camaldulensis ≠ ´
Myriophyllum aquaticum ´ ≠
Ambrosia confertiflora ≠ ´
Cyperus involucratus ≠ ´
Cyperus odoratus ≠ ´
51
Datura stramonium ≠ ´
Pennisetum clandestinum ≠ ´
Pistia stratiotes ´ ≠
Nicotiana glauca ≠ ´
Eichhornia crassipes ´ ≠
Xanthium strumarium ≠ ´
Ipomoea aquatica ´ ≠
Atriplex holocarpa ≠ ´
Salvinia molesta ´ ≠
Sesbania sesban ≠ ´
Ficus benghalensis ≠ ´
Ficus microcarpa ≠ ´
Ficus religiosa ≠ ´
Schinus terebinthifolius ≠ ´
Paspalum distichum ≠ ´
Ricinus communis ≠ ´
Populus alba ≠ ´
Acacia saligna ≠ ´
Acacia salicina ≠ ´
 can be found in these ecosystems and recent reports from the field seem to
confirm that the situation is getting worse as new foci of invasive plants are
regularly reported. The sensitivity of freshwater habitats and wetlands in Israel
is the result of a combination of several factors:
Fist, Israel has a dry season, i.e. a season with a negative water balance3, which
lasts for seven or eight continuous months, longer than found in any other
Mediterranean climate in the Mediterranean region (Dufour-Dror & Ertas 2004).
The natural seasonal drought is the most limiting abiotic factor for vegetation
cover in Israel. Yet, the constraint posed by this abiotic stress is significantly
smaller in wetlands and almost irrelevant in freshwater ecosystems. This can
partially explain why many invasive plants become successfully established on
stream banks, near winter pools, in seasonally flooded areas and in sites with a
high water table (figures 41a, 41b and 41c).
Secondly, as a result of the precipitation regime, which is concentrated in
winter months, streams, wadis and rivers in Israel are subject to strong
seasonal flooding. These floods act as highly efficient vectors of seeds and plant
52 fragments dispersal and therefore allow invasive plant species to disperse their
seeds over relatively long distances downstream and to spread in new areas
that are temporarily flooded.
However, these natural phenomena alone would not be sufficient for invasive
plants to take over moist habitats, had it not been combined with anthropogenic
activity that almost completely destroyed the natural vegetation cover along
streams and in wetlands, with the exception of protected areas. In many
cases natural vegetation was destroyed for landscaping reasons, or as part of
management treatments to prevent flooding, mainly in agriculture fields. Periodic
cutting of vegetation, mostly along rivers created a window of opportunity for
invasive plants to invade stream banks. On the other hand, in natural sites where
native species of riparian habitats were preserved, or restored, e.g. Oleander
(Nerium oleander), reed species (Arundo donax, Arundo mediteranea), or holly
bramble (Rubus sanguineus), the vegetation cover is literally impenetrable to
invasive species (figures 42 and 43).

The combination of these factors can explain why wetlands are among the most
vulnerable habitats to invasive alien plants in Israel.

2.3 – Managing invasive plants in Israel

The ultimate aim of the study of biological invasions is the preservation of
natural ecosystems and the conservation of biodiversity. The importance
of understanding the characteristics of invasive plants, their ecological
requirements, their biological traits and their spatiotemporal spreading patterns
are relevant mainly within the perspective of developing management strategies,
control methods and techniques that can be applied in the field as part of nature
conservation management programs.
3
A period in which the potential evapotranspiration (evaporation + transpiration) is higher than precipitation.
A

53

Figures 41. Three examples

of how invasive plant
species take over moist
habitats in Israel.
Figure 41a. Datura
stramonium on the banks
of Nahal Kidron, Dead Sea
Valley.
Figure 41b. Ambrosia
confertiflora on the banks
of the Alexander River in
the Sharon region.
Figure 41c. Xanthium
strumarium near a water
body in the Hula Nature
Reserve.

C
 Figure 42. Original
vegetation cover dominated
by Nerium oleander along
the banks of the Yavne’el
River in the eastern Lower
Galilee.

Figure 43. This photo was

taken in a dense stand
of Arundo mediterranea, a
native reed species planted
in experimental sites along
the Alexander River banks
54 to prevent the spread of
the invasive Ambrosia
confertiflora through the
rehabilitation of the natural
vegetation cover in this
type of habitat. The stands
created by this particular
species of reed have
so far been found to be
impenetrable to invasive
plant species.

The control methods that can be used in order to contain, reduce or prevent
the proliferation of invasive plants pertain to four categories: (1) physical or
mechanical techniques, (2) chemical treatments, (3) biological control, and (4)
preventive actions. As of now, the main methods applied in Israel are physical
and chemical. Classical biological control, based on the introduction of a natural
enemy of the targeted plant in local ecosystems is currently being discussed
in Israel but is not yet permitted. Preventive actions have been very recently
initiated by the Ministry of Environmental Protection, and are still in their very
first steps.

2.3.1 – Physical and chemical control methods applied in Israel:

an overview
Physical or mechanical methods
Different methods of physical or mechanical control have been experimented in
Israel in recent years. Physical control of invasives is currently applied locally in
a few Nature Reserves, as part of management programs under the supervision
of the Israel Nature & Parks Authority (INPA).
Figure 44. An individual of Acacia saligna resprouting only two weeks after cutting in the Judean
Mountains National Park.

In most cases physical or mechanical control of invasive plants in Israel is 55

carried out either by uprooting individuals, by cutting them down, or, in the case
of free floating aquatics, by simply collecting the plants from the water body.

Theoretically, cutting would have a number of advantages since it does not

involve the use of chemicals that could pollute the ecosystem, it is relatively
inexpensive, and it is generally easy to apply. Unfortunately, the efficiency of
cutting is very limited due to the high resilience and resprouting abilities of
almost all the invasive plant species found in Israel, which are able to recover
shortly after cutting. For example, when trees such as Acacia saligna or Acacia
salicina are cut down they quickly regenerate from the stumps and from their
root system (figures 8 and 44). The same is true for perennial herbaceous
species such as Ambrosia confertiflora whose vitality is not affected even by
repeated cutting. Other tree species, such as Ailanthus altissima, develop root
suckers that emerge a few meters from the stump and allow the tree to recover
very rapidly after being cut down (figure 45).

Figure 45. Sprouting and development of root suckers from a cut stump of Ailanthus altissima.
 Unlike cutting, uprooting is locally used by the INPA against the camphorweed
(Heterotheca subaxillaris) and the Hottentot fig (Carpobrotus edulis) in order to
reduce their densities in the sandy habitat of the Na’aman Delta Nature Reserve,
south of Acre in the Western Galilee (Sinai et al. 2011). In the same region
new foci of water lettuce (Pistia stratiotes) and the water hyacinth (Eichhornia
crassipes) were removed manually immediately following their emergence in
the Na’aman River (Ibid.) which prevented further local infestation.
However, uprooting does not necessarily ensure that the invasive species will
not renew itself, because in most cases some roots or root sections remain
in the ground and even a small root fragment can produce a new individual,
especially among taxa that are able to reproduce vegetatively. For this reason
the recent attempt to control Kikuyu grass (Pennisetum clandestinum) in the Ein
Afeq Nature Reserve in the Western Galilee, involved topsoil removal (Ibid.). In
addition, uprooting is feasible mainly for herbaceous species, and in some cases,
for dwarf shrubs or creeping species. Uprooting larger plants would disturb the
soil and enrich it with nitrogen, which in turn improves the soil conditions and
consequently facilitates the establishment of invasive leguminous tree species
56
such as Acacia and Parkinsonia species. Finally, uprooting does not affect the
seed bank in the soil from which entire population of invasives can reemerge
through the germination and development of new individuals.
In order to tackle this problem another type of mechanical control attempted in
Israel is solarization, which uses impermeable opaque plastic sheets to cover the
soil surface in order to induce seed germination. The high temperatures created
beneath the plastic cover eventually kill the young shoots. This control method,
whose purpose is to reduce the soil seed bank, was tried in Israel with Acacia
saligna seeds (Cohen et al. 2008). This technique is effective but it has many
drawbacks: its application is not selective and may harm local plant species, it
is relatively expensive, plastic sheets may be easily shred by wild animals and
it cannot be applied in all terrains since it requires a relatively flat topography.
Hence, this control technique is not implementable over large areas (Richardson
& Kulge 2008). Still, it may be applied locally in specific disturbed and semi
natural sites such as roadsides.
Another physical control method currently being tested in Israel is the
rehabilitation of natural riverbank vegetation in order to stop and prevent
the proliferation of the burr ragweed (Ambrosia confertiflora) along natural
watercourses. The first results of an experiment along the Alexander River
in the Sharon plain (Israel’s central region) that began in 2008 and is still in
progress, showed that only the stands formed by the native reed species Arundo
mediterranea, are impenetrable to the burr ragweed (Dufour-Dror 2010).
Physical or mechanical control methods have limited applicability in Israel and
for the reasons mentioned above they cannot be easily implemented over large
areas. Yet, in some cases, they can prove effective only on a very local scale,
as they may help to prevent the development of nascent foci of invasive plants
in protected areas.
Therefore, as the majority of invasive plant species in Israel cannot be
effectively controlled simply by physical and mechanical means, most of the
control actions currently implemented in Israel against invasive plants involve
chemical agents.

Chemical control methods

The use of herbicides against invasive plants in natural areas is the most
common control method in Israel. It should be noted that chemical control in
natural, and even protected, sites is not necessarily inappropriate provided it is
performed with care and according to specific procedures designed for control in
natural ecosystems (Wittenberg & Cock 2001, Burn 2003).
Chemical control of invasive plants in natural sites in Israel has been used
mainly against invasive trees. In 2005 a first experiment was conducted against
mature Acacia saligna trees and saplings in the Sorek River Nature Reserve
in the Judean hills (Dufour-Dror 2007a) and simultaneously against mature
individuals of Ailanthus altissima in open areas west of Jerusalem (Dufour-Dror
2007b). The targeted chemical control of these invasive trees was performed 57
using the drill-fill and the frilling techniques (see detailed description in Muyt
2001 and Dufour-Dror 2007a). The decision to use herbicide injection rather
than spraying was based on results of previous studies that showed that wattle
phyllodes are resistant to herbicides (McDermott, 1987, pers. comm., Pieterse
& McDermott, 1994). The herbicide used was glyphosate (undiluted) which is
considered to be of low toxicity to terrestrial fauna (Burn 2003). Since it was
directly injected into the stems of the invasive trees the risk of non-target
effect from this non-selective herbicide was eliminated (figure 46). The control
technique, based on drill-fill and frilling proved very effective: 83% of mature
Acacia saligna trees and 92% of mature Ailanthus altissima died after the first
treatment, as did more than 95% of the wattle saplings (Dufour-Dror 2007a,b;
figures 47 and 48). The key to the success of this simple control method, that
involves very small quantities of herbicide, is in the rigorous implementation of
the procedure, notably the injection of the herbicide into the xylem within ten
seconds after frilling.

Other experiments of chemical control against Acacia saligna were carried out
recently in the Einot Gibton Nature Reserve, a wetland located in the southern
coastal plain, heavily infested with golden wreath wattle, where a control
program was initiated by the Nature and Parks Authority. Chemical control was
performed with the cut-stump technique, initially with triclopyr and oil, but
many individuals were not fully eradicated and developed root suckers (Cohen
et al. 2010a). Triclopyr, which is very expensive, was replaced with fluroxypyr,
while oil, which may pollute the soil, was replaced with Silwet L-77, a surfactant
used to spray on phyllodes that developed on resprouting stumps (Cohen et al.
2010b). In this case the use of both fluroxypyr and Silwet L-77 in spray may
seem questionable considering the known adverse effects of these chemical
compounds in aquatic environments.
58

Figure 46. Three ways of applying targeted chemical control in woody species. Above:
drill-fill in Ailanthus altissima. Middle: frilling applied to Acacia saligna saplings. Below:
cut stump method applied to mature Acacia saligna trees
(bottom photo: Talia Oron).
The targeted chemical control method
using the drill-fill or the frilling
technique against invasive trees is
indisputably effective, inexpensive and
can easily be applied in all types of
terrains. Moreover control of the trees
in situ, i.e. with the drill-fill or the
frilling methods, significantly reduces
disturbances related to the removal
of cut trees. Nevertheless, this control
method is time costly as it requires
individual treatment of each invasive
tree. It therefore may be suitable for
controlling small and medium foci
but may be difficult to apply in large
populations.

Chemical control of herbaceous species
in natural areas in Israel is still rare
because of the non-target effect
concern. However, a water lettuce
infestation that choked a canal in
the Hula region (northeastern Israel)
was recently successfully controlled.
The relatively large population that
became established there in the early
Figure 47. Dead trees of Acacia saligna 59
following treatment by drill-fill, next to an
untreated tree in full bloom. Photographed
in the Sorek River Nature Reserve in 2006
when the targeted chemical control method
was tested in Israel. Follow-up during
several years showed that when the drill-fill
technique is applied properly the controlled
trees die within few months and do not
develop root suckers or sprouts, unlike the
results with the cut stump method.

2000s was controlled with ametryn, a

herbicide with low toxicity to terrestrial
and aquatic freshwater species
(Yaakoby, pers. comm. 2012). Even
though some water lettuce individuals
can still be found in the site, the bulk of
the population collapsed following the
treatment.

The question of chemical control was also

addressed recently for camphorweed
(Heterotheca subaxillaris) and a study
proved several herbicides are effective
(Quaye et al. 2010). Yet the chemical
control of this plant in natural areas
could pose serious problems of non-
target effect and further experiments
are needed in order to determine if
chemical control of this invasive in Figure 48. Dead young sapling of Acacia
natural sites should be considered. saligna a few weeks after frilling.
 2.3.2 – The issue of biological control against
invasive plants in Israel
The proliferation and the establishment of invasive plants over vast natural
areas have made physical and chemical control almost completely irrelevant, as
their implementation over large regions would be excessively time-consuming
and too costly. This is why biological control methods (biocontrol) have been
developed and applied against various invasive plants mainly in Australia, New
Zealand, South Africa and the United States. The basic principle of biocontrol
is to release a natural enemy of the targeted invasive plant in the infested
ecosystem. Therefore biocontrol is in effect the intentional introduction of
another alien species, usually an arthropod or a fungus, that is expected to pose
a threat on the targeted invasive so its proliferation will be hampered, or even
stopped. For example a rust fungus can cause irreparable damage to the foliage
of invasive plants while the larvae of some weevils feed on the invasive plant
seeds and destroy them even before they ripen (see figures 4 and 5).

The main advantages of biological control are the ease of application, the
60 relatively low cost, and mostly the fact that there is no need of repeated control
operations, i.e. if the introduction of the insect or the fungus is successful there
is no need for further action as the natural enemy released reproduces and
spreads in the area infested by the invasive plant. This last aspect is crucial in
extensive natural areas. Another advantage for managers is the rapidity of the
results: the effects of biocontrol are generally perceptible within few years after
implementation.

Many researchers, among them numerous entomologists, advocate biocontrol and

suggest it should be regarded as the most valuable control method for managing
alien invasive plants (Briese 2004; Palmer et al. 2010; Groenteman et al. 2011;
Moran & Hoffmann 2012). Indeed it seems that a large proportion of biological
control programs that targeted invasive plants in the countries mentioned above
achieved complete or partial control, and did not result in catastrophic events, to
the best of our knowledge at this point (Van Driesche et al. 2010).

However, there has been considerable debate on risks associated with biological
control as many researchers pointed out negative consequences that classic
biological control can have on natural ecosystems. One of the risks of biocontrol
is the non-target effect that occurs when the introduced natural enemy, which
is itself an alien species, begins to negatively impact native species (Brimner &
Boland 2003, Boland & Brimner 2004, Carson et al. 2008; Chalak et al. 2010,
2011; Simberloff 1996, 2012). The problem posed by the non-target effect has
led to better risk assessments that improved the ability to predict post-release
impacts (Barratt et al. 2010). Yet we still know little about the magnitude of non-
target effects (DeClercq et al. 2011). For instance several authors have recently
raised the question of the spread and proliferation of introduced weed biological
control agents to adjacent areas (Pratt & Center 2012; Simberloff 2012) an issue
which is probably very relevant on continents. The accumulation of bioagents
in local food webs also raises many questions since recent observations showed
the existence of unpredicted relationships between both local and alien species
with introduced bioagents (see in DeClercq et al. 2011).

The most recent observations about the rust fungus Uromycladium tepperianum,
introduced to South Africa to control Acacia saligna, which is also invasive
in Israel, provide an interesting illustration of the current questions and
uncertainties relating to the potential negative impacts of biological control as
mentioned above. The control of Acacia saligna with Uromycladium tepperianum
in South Africa has so far been an indisputable success (Morris 1997; Wood
& Morris 2007), even though it did not lead to eradication and requires
a combination with mechanical clearing and herbicide applications (Moran &
Hoffmann 2012). On the other hand non-target effects of the rust fungus have
been recently observed outside its natural range, in Indonesia and Malaysia,
with serious damage to local agriculture (Old & Dos Santos Cristovao 2003;
Rahayu et al. 2010). Moreover, interactions between another invasive, the
false codling moth (Thaumatotibia leucotreta) and the introduced rust fungus
have been observed in South Africa, where the alien moth uses the galls of the 61
introduced rust fungus as a larval food resource (Veldtman et al. 2011). The
issue of the proliferation of rust fungus to other areas of the world is also very
relevant, and the problems it may cause “cannot be anticipated at the present
time” (Wingfield et al. 2001).

More generally, it is very surprising to see that in the debate about the potential
effects of biocontrol, there is almost no mention of the fact that we may not
be fully able to assess the consequences of biocontrol simply because not
enough time has elapsed since most biocontrol projects began. About half of the
invasive plant biological projects worldwide are still in progress (Van Drieche et
al. 2010). We know that the dynamics of plant invasion require a lag phase that
may last several decades during which no serious impact on local ecosystems
can be noticed. So it would be reasonable to assume that most of the negative
consequences of biological control that may occur in natural ecosystems are yet
to be observed in the years to come.

To this day there is no biocontrol against weeds in Israel, as there is in Europe.

Yet the possibility and the need to introduce natural enemies of invasive plants
have very recently been suggested. However, the question of using bioagents
against the most problematic invasive plant species in Israel will have to be
addressed thoroughly. Before making the decision to release natural enemies
of invasive plants in Israel, it is essential to take the following facts into
consideration:

Although Israel has a Mediterranean and an arid climate its geographical

characteristics are radically different than these of South Africa or Australia
where many biological control programs have been implemented: Israel is a
very small country (22,000 km2), densely populated, with limited natural areas,
particularly in the Mediterranean region of the country. Moreover it is not an
 island, as it borders on four countries and it is located at the junction of Africa,
Asia and Mediterranean Europe. In this context it is not possible to ignore the risk
of a bioagent (an arthropod or a fungus) spreading to neighboring countries.

Moreover, the fact that biological control does not necessarily lead to eradication
of the invasive, as seen in South Africa with Acacia saligna and Uromycladium
tepperianum (Moran & Hoffmann 2012), signifies that other methods, such as
chemical control in the case of Acacia saligna, will have to be implemented if
we are to eradicate local foci of the invasive wattle. Since chemical control has
proved efficient when properly applied and considering the fact that infested
sites in Israel are restricted, incomparably smaller than the areas infested
in the fynbos of South Africa, the necessity of using biological control in the
case of Acacia saligna and other invasive wattles in Israel is questionable. The
small dimensions of infested areas in Israel, relative to the size of infested
areas in countries such as South Africa and Australia, suggest that treatment
with existing methods, including site-specific herbicide application, would be
preferable and definitely much safer for local ecosystems, despite their cost,
62 than biological control.

However, biological control in Israel should not be completely ruled out. It may
be the best option to control specific herbaceous invasive taxa such as the red
water fern (Azolla filliculoides) or the burr ragweed (Ambrosia confertiflora) that
are very likely to become the most problematic weeds in local ecosystems in
Israel, considering their very high spreading rate. The chemical control of these
invasive has considerable drawbacks as they invade wetlands. For very similar
reasons Ambrosia species, as well as free-floating invasive aquatics such as
Azolla filliculoides, are considered as good targets for classic biological control in
Europe (Gassmann et al. 2006; Gerber et al. 2011).

2.3.3 – Prevention of further plant invasions in Israel

Prevention is indisputably the most cost effective management measure among
all the actions that can be taken against invasive plants (Finnoff et al. 2006,
Hulme 2006). Several dozen alien plant species have already established
themselves in natural areas in Israel and the probability that additional species
will do so in the future remains very high. Therefore the most urgent step to
prevent further plant invasions is to avoid planting potentially invasive taxa that
have already been introduced to Israel and to prevent the introduction of plant
species not yet imported into the country. This is precisely the purpose of the
list of Israel’s least wanted alien plant species (IL-LWAPS) that was very recently
edited by the Israel Ministry of Environment together with the Israel Nature &
Parks Authority and the Israel Ministry of Agriculture (Dufour-Dror et al. 2012).
The list includes 143 taxa, all ornamental species, of which 47 (33%) have
not yet been introduced to Israel. Among the 96 species already introduced to
Israel, 48 (34%) have not yet escaped their cultivation areas, 15 (10%) are
casual, 10 (6%) are naturalized and 23 (16%) are already invasive.
The species included in the list were selected according to the approach that
assumes that a plant species which is invasive in a given area is very likely
to become invasive in other regions experiencing a similar climate as well
(Pheloung et al. 1999, Reichard 2001, Groves et al. 2001, Rejmanek et al.
2005). For instance it is highly probable that a plant which is invasive in the
Cape region, in Southeast Australia or in California, will also become invasive if
introduced to Israel’s Mediterranean region.

2.4 – Selection of species included in the book

The third section of this book provides detailed and up to date information on
a selection of 50 alien plant species found in natural areas in Israel. As the
objective of this book is to focus on the major invasive species and/or on the
taxa with a high invasion potential, the detailed information has been limited to
50 species. However a comprehensive list of all alien plant species recorded in
Israel’s natural and disturbed areas, compiled by Ori Fragman-Sapir from the
Jerusalem Botanical Gardens and the author appears at the end of the book. 63

The criteria for determining the 50 main alien plant species are very simple: Only
plant species invading natural areas were selected. Not included were species
found, for example, exclusively in built-up areas, roadsides or agricultural areas.
Species whose proliferation stage is “invasive” were included. To them were
added “casuals” or “naturalized” species that are known as major invasive taxa
in Mediterranean or arid areas, such as the black locust (Robinia pseudoacacia)
and the kangaroo thorn (Acacia paradoxa).

The invasive plant taxa listed in the third section of the book include species that
are almost unknown as invasive species outside Israel. These include Australian
wattles such as the bramble wattle
(Acacia victoriae) or the willow
wattle (Acacia salicina), but also
the camphorweed (Heterotheca
subaxillaris) and to a great
extent the pop saltbush (Atriplex
holocarpa) and the sacred fig
(Ficus religiosa).

A number of alien species whose

invasive status in Israel is still
unclear were not included.
For example, the limestone
wattle (Acacia sclerosperma,
figure 49) is a shrub that was
Figure 49. Acacia sclerosperma. This Australian
planted along roadsides in Israel’s
wattle was not included in the present edition of
Mediterranean region and on the the book since it is not yet clear if it should be
banks of wadis in the northern considered as an invasive species in Israel.
 Negev. Individuals that might have germinated and grown without human
intervention were spotted very rarely, but further observations are needed
in order to determine whether this species is “casual” or ”naturalized”. The
rosewood (Tipuana tipu) was not included among the 50 species for similar
reasons.

However, the dynamics of biological invasions are rapid and it is very likely that
in the near future additional invasive plants will join the list. It is even more
probable that some of the species whose invasive status is now defined as
“casual” or “naturalized”, will, in the near future, be defined as “invasive”.

64
65
Alien Invasive
Plants in Israel
 Acacia cyclops A.Cunn ex G.Don
Western Coastal Wattle
68
Family: Fabaceae (incl. Mimosaceae)
Basic Description: Acacia cyclops is a
low evergreen tree or shrub, 1-5 m tall.
It has alternate erect phyllodes, 4-9.5
cm long and 0.6-1.5 cm wide, with 3-4
prominent, parallel veins on their surface.
The tree has no spines. Its inflorescence
is a small raceme, 3-20 mm long, bearing
2 yellow globular heads. The heads are
composed of 60-75 minute flowers, and
its diameter is 5-7 mm. The pods are
brown, usually rounded with a slight
flattening between the seeds. The seeds
form clearly visible protrusions in the pod.
The pods remain on the branches for at
least a year after they open and release
their seeds. Another identifying feature is
the orange-red seed pedicel that envelops
each seed. Acacia cyclops blooms in Israel
from April to June.
Introduction to Israel: Acacia cyclops
was introduced by the British into Israel in
the 1920s for afforestation, in particular
soil stabilization.
Biological and Ecological Properties:
Acacia cyclops grows on most soil
types, including sands. It has a high
tolerance to drought and soil salinity.
It reproduces only from seeds, which
are produced in large numbers. South
African studies found that the Acacia
cyclops seed bank in the soil contains
between 2000 and 5900 seeds per square
meter. Unlike Acacia saligna, 95% of the
seeds germinate or die within one year.
In countries where Acacia cyclops is
invasive, e.g. South Africa, its seeds are
dispersed mainly by birds. In Israel it has
still not been determined which animals
are involved in dispersing Acacia cyclops
seeds. The main obstacle in eradicating
Acacia cyclops concentrations is the
extended seed viability in this species, 50
years. Fires facilitate seed germination
and intensify invasiveness. Acacia cyclops
grows rapidly, particularly during the
two years following germination, and its
lifespan is about 30 years. Acacia cyclops
has allelopathic properties.
Distribution in Israel: Acacia cyclops
possibly “escaped” from the JNF plant
nursery that once existed at Kiryat
Anavim in the Jerusalem Mountains
region. The main population in Israel,

which has hundreds of trees, is found
in the Judean Mountains National Park
where a huge wildfire occurred in the
summer of 1995. Lately it has been found
in a number of sites in open areas in
the Caesarea region (northern Sharon)
and south of Ashkelon in the southern
coastal plain. Should the tree become
established in the coastal sands along the
Mediterranean Sea shores, its preferred
habitat, it would probably proliferate
and cause serious ecological damage, as
it did along the coast in South Africa’s
Mediterranean region. Acacia cyclops has
been classified as “invasive” in Israel.
Effect on Native Species: Acacia
cyclops can form extremely dense stands
that displace native species by preventing
them access to light. Simultaneously the
leaves that fall from the tree accumulate
in a particularly thick leaf litter layer
that enriches the soil with nitrogen and
Geographic Origin: The natural range of Acacia
cyclops is southwestern Australia and along its
southern coast.
� References: 76, 149, 153, 155, 156, 164, 165, 182, 200, 212, 218, 247, 278, 287, 290, 340, 351.
encourages development of other invasive
plants that effectively use nitrogen,
such as other Australian acacia species.
In addition, unlike most other invasive
wattles in Israel, Acacia cyclops possesses
allelopathic properties: a study has
shown that its leaves release chemical
compounds that prevent germination of
other species.
Control: Unlike other Australian acacias,
cutting is relatively effective against
Acacia cyclops, but only if done close
to the ground. Apparently no chemical
control methods have been investigated
against Acacia cyclops. Biological control
methods have been developed and
implemented in South Africa: In 1991
a seed-feeding weevil (Melanterius
servulus), which is a natural enemy of 69
Acacia cyclops was released in the Cape
Region. The levels of seed destruction
measured from 1998 to 2009 averaged
only 56%. In 2001 a flower-galling fly
(Dasineura dielsi) was released in the
same region. The fly creates galls on
the acacia flowers and prevents pod
production. The results are variable: some
years the fly succeeds in preventing all
pod production in specific areas, but in
other years pods develop in spite of fly
activity.
Introduced Range: Acacia cyclops is one
of the most common Australian acacias in
South Africa. It is considered an invasive
species in the western United States and
has begun invading North Africa and
southern Europe.
 Acacia karroo Hayne
Karroothorn
70
Family: Fabaceae (incl. Mimosaceae)
Basic Description: Acacia karroo is
a shrub or tree; in Israel it is 5-6 m tall
(but can reach 25 m in South Africa).
Its leaves are 120 mm long and 5 mm
wide, pinnate with 8-20 leaflet pairs.
Ivory colored spine pairs, 10 cm long
and occasionally longer, are arranged
in a V-shape along the branches. The
inflorescence is a cluster bearing 4-
6 globular heads, 10-15 mm in diameter.
The yellow heads each contain some 90
minute flowers. Its pods are glabrous,
up to 16 cm long and 1 cm wide. They
are hooked and constricted between the
seeds. Acacia karroo blooms in Israel
from the beginning and throughout the
summer.
Introduction to Israel: Acacia
karroo was introduced into Israel as an
ornamental by the British between 1927
and 1930.
Biological and Ecological Properties:
Acacia karroo usually grows in areas with
over 500 mm annual precipitation. It is
tolerant of frost and hot weather and
can develop on all types of soil, including
sandy and saline soils. Its root system is
very dense and spreads both horizontally
and vertically deep into the ground.
Acacia karroo produces large quantities
of seeds: an adult plant can produce
up to 19,000 seeds yearly. In its natural
range the seeds are viable no longer
than one year. In Australia, however
seeds that penetrated the ground
preserve viability for up to 7 years.
Acacia karroo seeds are dispersed mainly
by birds. There is no information on
seed dispersal of invasive specimens in
Israel and on the extent of their variability
when dormant. This property indicates
a high degree of competitiveness
and ability to invade natural areas,
even undisturbed ones. Acacia karroo
regenerates from stumps after cutting or
fire. Its average lifespan is estimated at
about 40 years.
Distribution in Israel: Early signs
of Acacia karroo spread were found in
Israel on roadsides east of Caesarea (on
the western side of highway no. 4) in
the northern Sharon. A few trees were
found in the Ein Gev area located on the
eastern Lake Kineret shores, and in the
Hula Valley east of highway no. 90. In
2010 several individuals were found in
the Ramat Hanadiv protected site at the
southern end of the Carmel ridge. The
same year a few other individuals were
found in the Hula Nature Reserve in
the Hula Valley. The Acacia karroo
invasion in Israel is in its early stages
and at this time Acacia karroo is
classified as a “naturalized” species, but
its properties indicate that it has a high
invasive potential and poses threat
to most of Israel’s Mediterranean
region.
Control: There is still insufficient
knowledge on possible control methods
of Acacia karroo. Its ability to regenerate
following cutting makes it doubtful that 71
any type of mechanical control, including
fire can be effective. Infested sites in
Israel are still small, therefore it should
be still relatively easy to prevent further
proliferation throughout the country. The
effectiveness of targeted chemical control
as is applied against Acacia saligna and
Ailanthus altissima should be examined.
There is no biological control method yet
against this species.

Effect on Native Species: Acacia karroo Introduced Range: In recent years

can form very dense and impenetrable
units, which displace native species. Its
invasive ability could significantly reduce
the herbaceous species diversity in
infested sites. Since it is able to become
established in undisturbed natural areas,
new specimens are not always found soon
after their establishment and therefore
have time to reach maturity and produce
seeds.
Acacia karroo has invaded eastern,
southeastern and southwestern Australia,
where it is included in the National
Environmental Alert List of 28 new
invasive plants with the greatest invasive
potential. Acacia karroo grows in the
Mediterranean Basin in Libya, Morocco,
Spain, Portugal, Corsica and Sicily but has
not been classified as invasive in these
areas.

Geographic Origin: The natural range of Acacia

karroo extends throughout southern Africa and includes
southern Angola, Namibia, Botswana, Zimbabwe,
Mozambique, Lesotho, Swaziland and South Africa.

� References: 13, 101, 162, 231, 290.

 Acacia paradoxa DC.
Kangaroo Thorn, Prickly Wattle
72
Family: Fabaceae (incl. Mimosaceae)

Basic Description: Acacia paradoxa
is an evergreen shrub, 2-4 m tall, with
arched drooping branches. Its phyllodes
are narrow, 8-20 mm long and 2-7 mm
wide, oblong and end in a point. The
phyllodes are alternate, obliquely erect
and have undulated edges. At their base
are two spiny stipules, 4-12 mm long. The
golden yellow inflorescence is a solitary
globular head, 8-10 mm in diameter and
contains 30-50 minute flowers. Its fruit
is a straight or slightly bent pod, 6 cm
long and 3-5 mm wide. The pod is usually
covered with a dense layer of short white
hairs and has no constrictions between
the seeds. The seeds are black, glabrous
and oblong (3-5 mm). Acacia paradoxa
is considered especially polymorphic,
mainly in the shape of its leaves. It is
a particularly spiny acacia that blooms in
Israel in April.
Introduction in Israel: Acacia paradoxa
is believed to have been introduced into
Israel during the British Mandate as an
ornamental for gardens. It is not clear
whether it was used for sand stabilization,
as was the case in the United States. The
species was grown in the Kiryat Anavim
plant nursery in the Jerusalem Mountains,
which has not been in use since 1950.
Biological and Ecological Properties:
There is very little information relating
to the ecological properties of Acacia
paradoxa. A South African study from
late 2008 provided new information on
the species outside its natural range.
According to this study Acacia paradoxa
develops a vertical and horizontal
root system at the same time. This
provides the plant with effectiveness
and competitiveness in exploiting
groundwater, particularly in dry habitats.
Acacia paradoxa reproduces only by
seeds, which are dispersed, at least in
Australia, by ants. In the areas it invades
most of the seeds apparently fall near
the parent plant after the pods open.
Their dormancy period has not been
studied. Acacia paradoxa produces large
numbers of seeds, and in South Africa
concentrations of 1000 seeds per sq.
m were measured. The seedlings can
produce seeds as early as one year after
their germination. Acacia paradoxa is
tolerant of drought and cold, at least
down to -70C, and can grow in the shade
of other trees. In South Africa the tree
was found to regenerate following cutting.
In Australia its life span is estimated at
circa 20 years. It is not clear whether
Acacia paradoxa can survive for longer
periods of time in the area it invades.
Distribution in Israel: A few dozen
specimens of Acacia paradoxa were
found near the former JNF nursery, near
Kibbutz Kiryat Anavim in the Jerusalem
Mountains. They presumably “escaped”
and established themselves near the
road that passes nearby. Over the past
20 years a number of wildfires have
occurred in the area, which probably
accelerated the spread of this species in
the vicinity. Recent studies carried out in
South Africa suggest that Acacia paradoxa
has an extremely high invasive potential.
Therefore, there is a risk that this wattle
will continue to spread in Israel if the
present foci are not controlled. At this
point Acacia paradoxa is classified as a
“naturalized” species in Israel.
Geographic Origin: The natural range of Acacia
paradoxa is southeastern Australia.
� References: 82, 100, 149, 162, 200, 239, 279, 351, 361.
Effect on Native Species: Acacia
paradoxa can form dense stands that
displace native species. This is true for
moist habitats as well. In South Africa’s 73
Mediterranean region concentrations of 20
individuals per sq. m were measured.
Control: The evidence on the
effectiveness of mechanical control for
Acacia paradoxa is contradictory. In
Australia cutting is considered effective if
performed at the base of the trunk, near
the ground. In South Africa cutting was
found to be effective only when combined
with herbicide application (cut stump).
Spraying with triclopyr is only effective if
done during the growing season. There is
no method of biological control.
Introduced Range: Acacia paradoxa is
invasive in southwestern Australia, south
of Perth, and is spreading in recent years
as a result of fire. It is also invasive in the
south Western Cape Province in South
Africa, in lowlands along the southwestern
California coast and in northwestern New
Zealand along the coast.
 Acacia salicina Lindl.
Willow Wattle, Broughton Willow
74
Family: Fabaceae (incl. Mimosaceae)

Basic Description: Acacia salicina is
a large shrub, or low tree, 3-5 m tall
in Israel, with drooping branches. Its
phyllodes are pendent, with great
morphological diversity, narrowly linear or
slightly curved lanceolate, 7-20 cm long
and only 4-30 mm wide. The grayish-
green phyllodes are glabrous and leathery
with a single midvein. At the phyllode
base are 2-5 glands. The inflorescence is a
raceme 5-6.5 cm long, bearing 2-6 cream
colored or pale yellow globular heads.
The raceme has a diameter of 8 mm, and
bears 15-25 minute flowers. The pods are
long and narrow, up to 12 cm long and
7-13 mm wide, with constrictions between
the seeds. The bright black seeds are
oblong and 4.5-6mm long. Acacia salicina
blooms in Israel from late summer to
early December.
Introduction to Israel: Acacia salicina
was introduced into Israel either during
the British Mandate or after the State
of Israel was established. It has been
planted in Israel mainly in semi-arid
regions to stabilize stream banks and
probably also sands. Acacia salicina was
also planted in the Wadi Sekher area,
about 20 km south of Be’er Sheva in the
northern Negev, as part of agricultural
research that evaluated the species as
fodder in arid areas.
Biological and Ecological Properties:
Acacia salicina grows in areas with
more than 125 mm average annual
precipitation. It is resistant to drought
and cold up to -50C. Its preferred habitat
is clayish to sandy soils on stream banks
and flats in semi-arid regions up to the
edge of the arid region. Acacia salicina
is relatively tolerant to high soil salinity.
Its roots penetrate both vertically and
horizontally, and it is particularly fast-
growing, regenerating from stumps
after cutting. Acacia salicina reproduces
by seeds but also by root suckers. It is
relatively long-lived compared to other
invasive Australian acacias in Israel, and
its lifespan can reach 50 years.
Distribution in Israel: Acacia salicina
is invasive in the Negev along wadi
banks, particularly in its northern section
(Hatserim area, Agur Sands region
and central Negev anticline). A large
population has become established along
Wadi Shuval, northwest of Be’er Sheva
in the Hatserim region. A dense stand
developed on the eastern side of route
264. Other concentrations have become
established on most of the Northern
Negev wadi banks. Individuals developed
in the Negev Highlands Nature Reserve
and even north of the Yotvata Hai-Bar
Reserve located in the southern Arava,
in low areas where runoff concentrates.
A number of 5-m tall trees became
established at the mouth of Wadi Zohar
on the southern shore of the Dead Sea.
Acacia salicina is classified as “invasive”
in Israel.
Effect on Native Species: Acacia
salicina can form dense cover that
completely alters the character of local
vegetation cover, especially on wadi
banks. No other plant species grows
below its canopy. In addition to displacing
native species this shrub/tree competes
with native species for water in the soil.
The amount of biomass created by Acacia
salicina suggests that it consumes large
volumes of water, but this has not been
measured yet. 75
Control: Acacia salicina is not known
as an invasive species outside Israel,
thus there is no available information
on control methods for it. Its ability
to regenerate after cutting suggests
that mechanical control is not effective
against it. Several foci in the Negev were
controlled with the cut-stump method
but these were not monitored properly
so the efficiency of the method cannot
be evaluated. There is no method of
biological control, nor will any probably be
developed in the near future.
Introduced Range: Surprisingly, except
for Israel, Acacia salicina is not known
as an invasive species in other regions.
It has been reported as a naturalized
species in the Bahamas but has not been
classified as invasive there.

Geographic Origin: The natural range of Acacia

salicina is eastern Australia, except for the
northernmost tip of the area.

� References: 29, 72, 171, 182, 200, 203, 351.

 Acacia saligna (Labill.) H.L.Wendl.
Golden Wreath Wattle, Blue-leafed Wattle
76
Family: Fabaceae (incl. Mimosaceae)
Basic Description: Acacia saligna is
a shrub or low tree, 2-6 m tall, with
a single or multiple trunks, grey bark
and drooping branches. Its phyllodes are
very variable, dark green, glabrous and
leathery, linear or canaliculated, with
a prominent midvein. Both sides of the
phyllode are identical. Their surface is flat
or undulate, particularly the larger one.
The phyllode is 7-25 cm long and 4-20
mm wide and possess a distinctive gland
(1-2 mm) at its base. The inflorescence
is a raceme, up to 45 mm long, bearing
2-10 bright yellow globular heads, with a
diameter of 7-10 mm, each globular head
comprises 25-55 minute flowers. The
light brown pods are straight, with slight
constrictions between the seeds,
8-12 cm long and 4-6 mm wide. The
shiny black seeds are oblong and 5-6 mm
long. Recent genetic research has shown
that 3 to 4 separate subspecies of Acacia
saligna can be differentiated in Western
Australia. However, their morphological
taxonomic classification is complicated
and has not yet been formally recognized.
At this stage the informal subspecies are
subsp. lindleyi, subsp. pruinescens, subsp.
saligna and subsp. stolonifera. According
to a study currently being conducted in
South Africa, nuclear genetic materialis
shared between Israeli populations and
all the native genetic groups of Acacia
saligna (Thompson pers comm.). Acacia
saligna blooms in Israel from the end of
March to the beginning of May.
Introduction to Israel: Acacia saligna
was introduced to Israel in the early years
of the British Mandate for the purpose
of stabilizing sands, afforestation and
landscaping. The species was also planted
in Cyprus by the British during the same
period. Since its introduction, Acacia
saligna has become invasive in both
countries, but is absent from Lebanon and
Syria that were not under British rule.
Acacia saligna continued to be planted
in Israel at least until the 1980s as an
ornamental and for afforestation. In the
mid 1990s experiments were conducted
to examine the suitability of Acacia
saligna as fodder for sheep and goats in
desert areas.
Biological and Ecological Properties:
Acacia saligna grows in areas with
Mediterranean climates. It penetrates the
edges of semi-arid areas down to the 240-
mm isohyet, but does not develop in more
arid areas. Acacia saligna grows in average
annual temperatures between 110C and
230C. It can survive freezing temperatures
but temperatures below -60C are fatal. In
its natural range the tree grows in most
types of soils, including sandy soils and is
resistant to high soil salinity levels (up to
10-20 dS/m). The trees develop horizontal
roots up to 12 m long as well as vertical
roots that reach depths of 3.5 m, and up
to 16 m in sandy habitats. Reproduction
is both vegetative and by seeds. Two-year
old saplings can already produce seeds,
but most of the seeds are produced by
trees six years and older. Despite less than
1% of the flowers producing seeds, each
tree still produces very large numbers of
seeds. In South Africa trees produce on
average 5500 seeds/m2/year, and result
in very large, accumulated seed banks.
However up to 46,000 seeds/m2 have
been recorded for dense infestations.
A recent study in South Africa has shown
that a major portion of the seed bank is
situated in the upper 10 cm of the soil
profile. According to Australian studies up
to 85% of the seeds in the soil are viable
and are able to remain viable for extremely
lengthy periods – over 50 years. A South
African study found that local ant species
(Pheidole spp. and Anoplolepis spp.)
disperse Acacia saligna seeds to distances
of 2-3 m and bury them 4-7 cm in the
ground, thus reducing the exposure of 77
seeds to fire. Seed dispersal in Israel has
not yet been thoroughly studied but it is
likely that a common local ant species,
Pheidole pallidula, which also feeds on
the seeds, plays a significant role in
dispersing Acacia saligna seeds. Exposure
to fire triggers germination. According
to research carried out in Australia,
subspecies stolonifera and pruinescens
produce many root suckers; while
subspecies lindleyi produces relatively
few root suckers, while subspecies saligna
develops root suckers on rare occasions
only. All the subspecies can regenerate
from stumps after cutting and fire. Acacia
saligna has a particularly short lifespan,
usually 5-12 years and sometimes up
to 20 years in its native range. In Chile,
30-year old healthy specimens were
found at the peak of their development.
It is possible that the lifespan of the tree
increases significantly in areas where
it is invasive. A large population that
established after a large fire that occurred
in 1995 in the Judean Mountains National
Park (Jerusalem Mountains), showed first
signs of senescence in late 2010, i.e.
15 years after the fire event. However,
no studies have been conducted on the
lifespan of invasive Acacia saligna in
Israel.

Distribution in Israel: Acacia saligna is
currently the most notorious invasive plant
in Israel. With the exception of the Golan
78 Heights, the species is invasive throughout
the entire Mediterranean region in Israel;
from sea level up to altitudes of about
700 m. Acacia saligna creates very dense
units in disturbed habitats, particularly
on roadsides, but it also invades a wide
variety of natural habitats such as slopes
in the mountainous regions and the coastal
sands on the Mediterranean seashore
where it forms the densest stands. Many
protected areas are infested by very large
populations of Acacia saligna, e.g. the
Poleg Stream Reserve in the northern
Sharon, the Nitsanim Sands, Palmahim
Sands, and Einot Gibton reserves in the
southern coastal plain, the Bay Sands
Reserve in the Galilee coastal plain, and
Sorek River Reserve in the Jerusalem
Mountains area. The establishment of the
large population in the Judean Mountains
National Park is a direct result of the
large fire that occurred in 1995. Although
introduced in the 1920s A. saligna was
designated an invasive species in Israel
only in the late 1970s, suggesting it
requires a minimum residence time of
about 50 years. Acacia saligna has been
classified as an “invasive” species in Israel.
Effect on Native Species: Acacia saligna
affects native species and ecosystems in
various ways: the dense, closed vegetation
units displace native species, as most
of them cannot survive in the shade
of the trees. Simultaneously, the thick
layer of dead phyllodes that accumulate
beneath the trees can in some cases
prevent germination of native species.
Moreover, this large amount of phyllode
litter enriches the upper soil layers with
nitrogen, thereby modifying the local
nutrient balance. This change facilitates
the establishment of other invasive plant
species that favor high soil nitrogen
concentrations. It has been shown that
the soil nitrogen level remains above
normal for a number of years even
after Acacia saligna has been removed,
facilitating invasion and establishment
of other alien invasive species. Acacia
saligna invasions in sandy habitats
substantially modify the structure of
vegetation cover and consequently
the character of these habitats. The
formation of a dense cover of trees
instead of an open, discontinuous,
dwarf shrubs and herbaceous cover is
presumably a significant change for the
psammophilic fauna that is accustomed
to open habitats. This change is even
more extreme in sites where Acacia
saligna is accompanied by Heterotheca
subaxillaris, e.g. the Bay Sands Nature
Reserve. Future studies will have to
quantify this effect in sandy habitats in
Israel. The establishment of dense stands
of Acacia saligna in dry habitats such
as sands or mountain slopes inevitably
reduces the amount of available soil
moisture for native species, particularly
woody natives. This effect has not yet
been quantified and studies are needed to
evaluate the effect Acacia saligna has on
the local water balance in infested areas,
particularly in sandy habitats. Another
negative effect of the invasion of Acacia
saligna on local ecosystems is related to
the fact that the biomass of Acacia saligna
trees potentially increases fuel loads and
consequently the intensity of fire events.
A recent review of the impacts of invasive
Australian acacias showed how a fire
event in an infested area depleted native
seed banks and ensured the domination
of Acacia saligna. The expected increase
in wildfire frequency in the coming years
in Israel, mainly due to the increasing
population density and development,
may therefore worsen the proliferation
of Acacia saligna in natural areas.
Acacia saligna is considered an “invasive
transformer” in Israel.
Control: Mechanical control options for
Acacia saligna are very limited, as the
species regenerates following cutting or
fire, thereby rendering these methods
ineffective. Uprooting is also ineffective
as some root fragments remain in the
soil and enable regeneration to take
place. There have been attempts to use
solarization to destroy seed banks. This
method proved effective for depleting the
seed bank in experimental plots but it is
not a viable option for heavily infested
sites, because the plastic sheets are
easily removed or damaged. An effective
method of mechanical control is covering
the ground with a layer of soil, at least 20
cm thick. This is however only effective
for landscape rehabilitation implemented
following infrastructure work, and not
for natural areas. Following a number
of studies conducted in South Africa
to assess control methods for Acacia
saligna, experts determined that the
most effective strategy is to reduce and
neutralize seed sources, i.e. destroying
all trees over the age of 5 years. Targeted
chemical control, using the drill-fill
technique for mature trees and the filling
method for saplings and seedlings has
been tested experimentally in Israel
and was successful. Targeted chemical
control with minute amounts of undiluted
glyphosate kills mature and young trees
at rates of 80% or more with no need
for repeated applications. This is a very
effective method particularly suitable
for implementation in natural protected
areas. It is easy to apply, it does not
involve cutting, which stimulates the
development of root suckers, and it is
inexpensive. This method however, is
time consuming and consequently suited
mainly for small and medium-sized foci.
South African scientists have developed
a biological control method with a rust
fungus (Uromycladium tepperianum)
imported from Australia and released in
1987 in the South African fynbos. The
introduction of the fungal pathogen
succeeded in reducing the tree density
in infested areas by 87-98%. Monitoring
showed that the spread of the fungus
reduced the lifespan of most of the trees
79
to about 7 years. Despite the promising
results, this treatment alone cannot
eradicate the Acacia saligna invasions as
the large seed banks are still problematic
as they provide a continuous source of
new propagules. In 2001 in South Africa
the rust fungal agent was supplemented
with a seed-feeding weevil (Melanterius
compactus) that destroys over 90% of
seeds while they are still in the pods on
the tree. In Israel no biological control
method against Acacia saligna has
been implemented but the possibility is
currently being assessed.
Introduced Range: Acacia saligna is
the most damaging invasive species in
the coastal lowlands of the south Western
Cape in South Africa. The species is also
invasive in Chile, Spain, Portugal and
Cyprus. In North Africa the tree has been
cultivated for 100 years, but has not
become invasive in the region, probably
due to the arid climate in the area.

Geographic Origin: Acacia saligna is endemic to

southwestern Western Australia. Most of its natural
range is located south of the 300-mm precipitation
isohyet (equal precipitation line).

� References: 52, 71, 72, 83, 114, 123, 153, 154, 155 , 156, 165, 182, 186, 200, 202, 205, 210, 211,
212, 218, 219, 229, 232, 272, 273, 278, 289, 302, 318, 325, 325a, 328, 335, 351, 353, 357, 358.
 Acacia victoriae Benth.
Elegant Wattle, Bramble Wattle
80
Family: Fabaceae (incl. Mimosaceae)
Basic Description: Acacia victoriae is
a shrub or tree, 2-5 m tall, occasionally
reaching 9 m. The greenish to grey
phyllodes are extremely variable in shape
and size. They are linear, or narrowly
elliptical, straight or slightly curved (2-5
cm long, 2-8 cm wide). Both stipules at
the phyllode base are spines, 2-12 mm
long. The inflorescence is a raceme up
to 10 cm long, bearing 15-30 yellowish-
cream globular heads. Each head is
composed of 15-30 minute flowers. The
light brown pods are oblong and relatively
flat, 8 cm long and 9-16 mm wide. The
seeds lie horizontally in the pod and form
clearly visible protrusions in the pod.
The seeds are dark brown, 4-6 mm long.
Acacia victoriae blooms in Israel from
mid-March to the beginning of May.
Introduction to Israel: Acacia victoriae
was introduced for afforestation,
apparently after the State of Israel was
established. It was planted mainly in the
northern Negev. The first invasive site
was discovered in late 2005 between the
Kama junction and the city of Rahat in the
southern Judean foothills.
Biological and Ecological Properties:
Acacia victoriae is particularly drought
tolerant, apparently thanks to its root
system that penetrates the soil to a depth
of about 20 m. Its preferred habitats are
beds and banks of wadis in arid and semi-
arid regions. Acacia victoriae grows rapidly
and in its natural range has a lifespan of
only 10-15 years. It produces particularly
great numbers of seeds and at the
beginning of summer bears an abundance
of pods. The tree grows in diverse soils,
but mainly in heavy soils on flats or
alluvial soils. Acacia victoriae regenerates
rapidly following cutting or fire, from
stumps and by root suckers.
Distribution in Israel: Acacia victoriae
is invasive in the Northern Negev,
particularly in the extensive planted
areas north of Be’er Sheva (east of route
40) in the southern Judean foothills. In
this area, which was planted in the late
1980s and early 1990s, thousands of
individuals germinated and grew alongside
the planted trees. At the site were Acacia
victoriae was first found invading in 2005,
east of route 264 (south of the Kamah
junction), a population of thousands has
developed forming a dense thicket. Other
invasive sites exist south of Latrun (on
both sides of route 3) in the northern
Judean foothills. Lately specimens began
spreading along route 1, from the Latrun
junction eastward. Others germinated and
grew in Levi Eshkol forest near the Mt.
Yavne’el Nature Reserve, southwest of Lake
Kinneret in the northern Jordan Valley. The
invasive scope in the Negev is reminiscent
of the Acacia saligna invasion along the
Coastal Plain. Although Acacia victoriae is
invasive in Israel for less than 10 years the
rapidity of its spread and the scope of the
infested areas justify classifying it as an
“invasive” species in Israel.
Effect on Native Species: Acacia victoriae
is capable of forming dense, occasionally
impenetrable thickets that displace native
species. Its water consumption has not yet
been studied, but it is likely that Acacia
victoriae concentrations use significant
amounts of water at the expense of native
plants. The vegetal unit it creates in arid
and semi-arid areas exhibits a forest-like
Geographic Origin: The natural range of Acacia
victoriae is throughout Australia, particularly in arid and
tropical areas.
� References: 20, 182, 200, 201, 351.
structure that is not suited for the desert
fauna. This may attract and encourage
nesting of non-local avian species such as 81
the hooded crow (Corvus corone cornix)
that can have a negative effect on the
population stability of local bird species.
Control: Acacia victoriae is not known
in the world as an invasive plant species,
therefore there is no extant information
on control methods. Cutting is not
effective because the tree regenerates
from root suckers. There is no information
at this time regarding its susceptibility
to herbicides and there is no available
biological control method.
Introduced Range: So far no reports
of Acacia victoriae invasions have been
recorded in the world, except in Israel.
Possibly the species was used for
afforestation mainly in Israel and is thus
unknown as an invasive species in other
arid and Mediterranean climate zones in
the world. In Australia, where it is native,
it sometimes becomes a nuisance around
watering points.
 Ailanthus altissima (Mill.) Swingle
Tree-of-Heaven, Chineese Sumac
82
Family: Pontederiaceae
Basic Description: A winter deciduous
tree that in Israel can reach a height
of 20 m. It has a straight grey trunk,
which is smooth in young specimens and
develops fissures in mature trees. Trees
that have been cut or burned develop
multiple trunks. Ailanthus altissima has
pinnate leaves 30-90 cm long, with 10-
40 lanceolate to oblong leaflets, 3-14 cm
long and 25-50 mm wide. The undersides
of most leaflets have 2-4 glands that
exude a bad smell. Ailanthus altissima is
dioecious, its small flowers are white or
greenish and arranged in panicles about
60 cm long. Its yellowish fruit, which
gradually turns light brown, has broad
wings, 4 cm long and 1.5 cm across.
Each fruit contains a single seed. In
Israel it blooms from May through early
June.
Introduction to Israel: Ailanthus
altissima is an ornamental tree that
was already common in Israeli gardens
in the early 20th century. The species
was planted in Jerusalem streets and
boulevards, particularly during the 1960s
and up to the 1980s.
Biological and Ecological Properties:
Ailanthus altissima is a fast-growing tree,
particularly in the first years of its life.
Young trees in Israel can grow at the rate
of a meter and a half a year. The tree
grows in areas that receive over 500 mm
of annual precipitation. Ailanthus altissima
is resistant to seasonal drought thanks to
its root system that is well developed both
vertically and horizontally. The species
can withstand cold temperatures as low
as -200C. It grows in most types of soils,
both acidic (below pH = 4.1) and basic,
shallow and deep, including sandy and
saline soils. The species reproduces via
seeds, but also via root suckers that can
reach a distance of some 15 m from the
trunk. Seeds are formed as early as three
to five years following germination. One
adult can produce up to one million seeds
annually that are dispersed by wind and
water. A new study found that immersion
in water improves the germination rate.
In California and Australia the seeds are
dispersed by birds as well, but this has
not yet been determined in Israel. The
seeds are viable in the ground for only
one year. Ailanthus altissima exhibits

distinct allelopathic properties: its
leaves release chemicals that prevent
germination of native species within a
5-m radius around the tree. Its average
life span is about 100 years.
Distribution in Israel: Ailanthus
altissima invades mainly open areas
and roadsides in the mountainous
Mediterranean region in Israel. A large
population established itself west of
Jerusalem in the Nahal Halilim Nature
Reserve and its surroundings. In northern
Israel there are a number of populations
in protected areas such as the Mt. Tabor
Reserve and the Tsipori National Park in
the central lower Galilee, and the En Afek
Reserve in the Galilee coastal plain. The
tree is also invasive in the Mt. Carmel
National Park and in the Ramot Menashe
area. Recently specimens have also been
observed outside the mountainous area,
near Modi’in in the northern Judean
foothills. Individuals have also been
observed lately on sandy loams in the
northern Sharon. It is estimated that
without immediate implementation of a
comprehensive control program Ailanthus
altissima will invade additional areas,
including so far undisturbed natural areas.
Geographic Origin: The natural range of Ailanthus
altissima extends from Eastern and southern China to
North Vietnam.
� References: 22, 32, 56, 84, 85, 131, 162, 180, 181, 185, 197, 207, 221, 239, 289, 302, 340.
The status of the species in Israel is
“invasive”.
Effect on Native Species: Ailanthus
altissima forms dense stands that displace
most of the woody native species due
to the heavy seasonal shade created
by its dense foliage. Its allellopathic
properties prevent native species from
germinating in infested areas. The species
is competitive thanks to well-developed
root system that utilizes soil water and
nutrients efficiently. Ailanthus altissima is
very allergenic, particularly to people who
suffer from asthma.
Control: Mechanical treatments, including
cutting and burning are completely
ineffective against Ailanthus altissima due
to its ability to regenerate from stumps.
Moreover, mechanical treatments such 83
as cutting induce the development of
numerous root suckers. Targeted chemical
control was found to be effective. A study
conducted in Israel in 2005 (whose results
were confirmed by a subsequent Canadian
study) showed that drill-fill treatment
with undiluted glyphosate (Roundup)
destroys over 90% of the trees. Grazing
in sites with young plants could suppress
Ailanthus altissima populations and serve
as an interim solution. No biological
control method has been developed yet.
Introduced Range: Ailanthus altissima
is a powerful invasive in France,
Switzerland, in southern and eastern
Europe, in southeastern and southwestern
Australia, in Canada, throughout the
United States (particularly California) and
in the Azores Islands. The species is also
invasive in North Africa, the southern and
eastern regions of South Africa, in New
Zealand, Mexico, Chile, Argentina and
Hawaii.
 Ambrosia confertiflora DC
Burr Ragweed, Slimleaf Bursage
84
Family: Compositae (Asteraceae)

Basic Description: An upright perennial introduced into Israel. It is possible that

herb, 75-250 cm tall. Its grayish green seeds were introduced in seed mixes for
leaves are 12-16 cm long and 10- feeding birds and pond fish, imported
15 cm across. The leaves have short from the United States.
petioles and are bipinnate, divided into
Biological and Ecological Properties:
long lobes; their margins have sparse
Ambrosia confertiflora generally forms
short hairs. The leaves are opposite at
very dense stands and is an extremely
the base of the stem and alternate on
competitive plant. It has a shallow, very
the upper part of the plant. Ambrosia
dense root system concentrated in the
confertiflora is monoecious and has male
upper soil layers extending down to a
and female yellow or greenish flowers.
depth of only 30 cm. It reproduces from
The male flowers are numerous and
seeds as well as vegetatively by adaptive
small (diameter about 1 cm) and are
buds found on spreading horizontal
borne on erect clusters. The female
roots. The prevalent reproduction mode
flowers lack petals and are concentrated
of the plant in Israel is still unclear. The
in the leaf axils in a cup-shaped
fruit is a bur with 10-20 hooked spines
involucre. Its seeds are brown and their
enveloping a single seed. The spines
diameter is 3-4 mm. In Israel the species
stick to the coats of animals and thus the
blooms in September-October
seeds are dispersed. Dispersal can also
occur by flowing water, particularly during
Introduction to Israel: Ambrosia
floods. The plants renew themselves
confertiflora was first recorded in Israel
within five weeks of mowing. Low winter
on the Carmel coast in 1990. Between
temperatures in Israel do not affect the
the late 1990s and the early 2000s,
plant survival.
a massive invasion began along the
Alexander River bank. There is no Distribution in Israel: The most recent
information on how the species was survey established that the infestation
began from the city of Shekhem (Nablus)
in Samaria from where the plant spread
first to the west along the banks of the
Shekhem Stream down to the banks of
Alexander River. A survey conducted
by the Sharon Drainage Authority at
the end of 2004 found that Ambrosia
confertiflora invaded 480 ha along the
banks of these two rivers and from there
it penetrated nearby fields and orchards.
The plant also began spreading eastward
along Wadi Tirtsa in eastern Samaria
where it forms very large stands, and it
recently reached the Jordan valley, north
of Jericho. At the end of 2008 Ambrosia
confertiflora was reported on the banks of
the Wadi Kaneh in the Wadi Kaneh Nature
Reserve located in the western Samaria
Mountains. Since 2009 several new small 85
foci were reported in the northern and
southern Sharon regions as well as on
the southern coastal plain. In March 2010
two foci were found along Route no.
1 in the Jerusalem Mountains. In spring
2011 the plant was reported for the first
time in the Galilee coastal plain, north of
Haifa. Ambrosia confertiflora is the fastest
spreading terrestrial invasive species in
Israel and may become a regional plague.
The Ministry of Environmental Protection
 is currently supervising the preparation
of a national plan in order to prevent the
86 spreading of the species in Galilee. As
Ambrosia confertiflora began spreading
about 15 years ago, it can be classified
as an “invasive” species in Israel.

Effect on Native Species: Ambrosia

confertiflora forms very dense units
that displace all other plant species, so
that the understory at infested sites is
completely lacking native species. The
plant radically modifies the vegetation
cover and consequently the whole
ecosystem. Ambrosia confertiflora is
a serious nuisance in cultivated fields
and citrus groves. It is not yet clear if
Ambrosia confertiflora has allergenic
properties similar to these of other
Ambrosia species.

Control: Mechanical control is useless

and can even exacerbate the intensity
of the invasion. Chemical control has
been implemented in Australia and
the herbicide that was found efficient
is Tordon 75-D which is composed
of 2,4-D and picloram. However, the
use of picloram in humid habitats
may have adverse effects and the
possibility of using this herbicide in
Israel has been rejected. Experiments
with other herbicides are planned and
will be carried out in the course of
2012. Another study, conducted along
the Alexander River is examining the
possibility of mitigating Ambrosia
confertiflora invasion by rehabilitating 87
native riverbank vegetation, chiefly with
the native reed Arundo mediterranea.
The experiment began in 2008 and
is scheduled to end in summer 2012.
Initial results showed that dense stands
of the native Arundo mediterranea are
impenetrable to Ambrosia confertiflora.
The species is invasive in very few
areas in the world so that no biological
control methods have been developed
for it so far. Nevertheless, following the
report, in 2008, of Epiblema strenuana, a
stem-galling moth found on specimens
of Ambrosia confertiflora in Israel, the
possibility of controlling the plant with
its natural enemy, accidently introduced
to Israel, is currently under study in the
country.
Introduced Range: Ambrosia
confertiflora is invasive in Western
Australia since the 1950s.

Geographic Origin: The natural range of Ambrosia

confertiflora is the semi-arid valleys in the southern
USA and Mexico.

� References: 10, 64, 86, 239, 354, 355

 Atriplex holocarpa F. Muell.
Pop Saltbush
88
Family: Chenopodiaceae
Basic Description: Atriplex holocarpa
is an annual plant 15-25 cm tall, with
alternate, succulent and rhomboid leaves.
The leaf margin has small, irregular
lobes, and the leaf is 10-30 mm wide.
Leaf color is greenish with white spots.
Atriplex holocarpa is monoecious, has tiny
separate male and female flowers that
develop in the upper leaf axils. The fruit
is spherical, lemon-like and has a spongy
texture. In Israel Atriplex holocarpa
blooms from June to December.
Introduction to in Israel: Atriplex
holocarpa was introduced into Israel in
the early 1960s to improve forage in
Israel’s arid and semi-arid regions. It
was planted in a number of experimental
plots in the Be’er Sheva area, northern
Negev, in the mid 1960s but did not meet
expectations and the experiment was
stopped.
Biological and Ecological Properties:
There is very little information on
the ecological properties of Atriplex
holocarpa. It grows in clay soils, in flat
areas and areas where runoff collects,
such as roadsides and wadi beds in the
desert. It does not grow in sandy soils.
Atriplex holocarpa is considered very
resistant to soil salinity and in Australia
is common along saline lake shores. The
species is wind-pollinated and its seeds
are dispersed by wind and runoff. There
is no information on how long Atriplex
holocarpa seeds remain viable in the soil
or whether wildlife feed on it.
Distribution in Israel: In Israel the
pop saltbush was first observed in open
spaces in the Northern Negev, i.e. a
number of years after it had been planted
in the experimental plots in the region.
Since then it has invaded roadsides in the
Hatserim area and in the central Negev
anticline region. The species continues
to spread towards the eastern Northern
Negev, e.g. Scorpion’s Ascent and it has
reached the northern Arava and the the
Dead Sea shores. Atriplex holocarpa
grows at the mouth of Wadi Tse’elim
on the Dead Sea shore and along Wadi
Darga in the southern Judean Desert. In
the Dead Sea Valley it grows mainly on
the roadsides of Route no 90 and a large
population was recently reported east of
Jericho. It has also been recorded in the
central Negev. The species is obviously
invading Israel’s arid region rapidly and is
classified as “invasive”.
of Atriplex holocarpa invasion on native
species.
89
Control: There is no information on
control methods of Atriplex holocarpa.

Effect on Native Species: The effect Introduced Range: At this time, in

of Atriplex holocarpa seems to be limited
to competition with other herbaceous
species for soil water, although it has not
yet been empirically quantified. Future
studies should assess other consequences
addition to Israel’s arid region, Atriplex
holocarpa is known to be invasive only
in Texas. There too the species began to
spread following attempts to use it for
forage.

Geographic Origin: The natural range of the pop

saltbush is the semi-arid areas in Southern Australia.

� References: 61, 104, 116, 217, 301, 302, 317.

 Azolla filiculoides Lam.
Red Water Fern, Large Mosquito Fern
90
Family: Azollacea
Basic Description: Azolla filiculoides is a
perennial floating aquatic fern, 25-50 mm
long. It has green or reddish overlapping,
scale-like leaves, only 1-1.5 mm long.
The plant has a short rhizome, from which
short, thin roots descend, floating in the
water. When the plants are exposed to
strong sunlight or low temperatures the
leaves produce red pigments.
Introduction to Israel: Azolla
filiculoides was introduced into Israel
as an ornamental aquarium plant at an
unknown time.
Biological and Ecological Properties:
Azolla filiculoides, like other species
belonging to the genus Azolla has
a mutualistic relation with the
cyanobacterium Anabaena azollae that
grows in pockets on the underside of the
plant’s leaves. The cyanobacterium fixes
atmospheric nitrogen, thus enabling
A. filiculoides to grow even in water
with low nitrogen concentrations. The
preferred habitat of Azolla filiculoides is
sluggish water bodies. Its populations
barely become stable in running water,
even when the flow is not fast. Azolla
filiculoides has a very rapid growth rate:
at water temperatures of 15-200C it can
double its surface coverage within 7 to
10 days. The fern dies at temperatures
below -40C, although specimens trapped
in ice can survive temperatures as low
as -150C. Azolla filiculoides reproduces
vegetatively as long as the water surface
is not completely covered. Once the
water surface is completely covered the
ferns reproduce by spores released in the
water. The spores allow the population
to recover and renew itself, even when
all the floating fern biomass has died,
which can be seen at the end of the
summer. Accumulating spores in the
water is usually the way Azolla filiculoides
populations can survive the winter low
temperatures.
Distribution in Israel: Azolla filiculoides
was first found in Israel in the mid-1980s
at the Dora Pool in the northern Sharon
near the city of Netanya, and later in that
same decade in the Bar-On Pool Reserve
in the northern Golan Heights. A large
population of Azolla filiculoides was found

in the Yarkon River, north of Tel-Aviv, in
July 2009, in the boundaries of the Yarkon
and Tel Afek National Park, at a site where
the water flow is very slow. The population
spread over 1⁄4 hectare together with
numerous specimens of Salvinia molesta
and a few specimens of Pistia stratiotes.
More recently, in May 2010, the fern was
found in the seasonal pond of the Birkat
Ya’ar Nature Reserve, northern Sharon,
where it formed the largest infestation
ever reported in Israel (5 to 7 ha). Its
recent proliferation in the country has
probably been caused by emptying
aquarium contents into natural water
bodies. Due to lack of regular monitoring
of populations in Israel, at this point in
time Azolla filiculoides can be classified
only as a “casual” species. Following the
recent infestation in the Sharon region
there is great concern that additional water
bodies will be affected in the near future
since small fragments of the fern may be
dispersed by water birds to new ponds and
water bodies.
Effect on Native Species: The formation
of a thick, impervious mat on the water
Geographic Origin: Azolla filiculoides has an extensive
range that includes South and Central America, Mexico,
the southwestern United States, western British
Columbia in Canada and the southern part of Alaska.
� References: 44, 100, 104, 106, 134, 149, 151, 162, 163, 263, 332, 340.
surface very significantly modifies
the abiotic conditions of the aquatic
ecosystem; therefore Azolla filiculoides is
considered a “habitat transformer” species
in the areas it invades. The plant reduces
the concentration of oxygen in the
water and lowers the pH. These changes
have a negative effect on most aquatic
organisms so that a continuous invasion
of Azolla filiculoides can negatively impact
biodiversity in the infested water body.
Azolla filiculoides invasions also interfere
with water birds that require open water
for their activity.
Control: In very small foci removing
plants with fine meshed nets may be
an option. It is advisable to use floating
barriers as well. However, since the
ferns are very small and disperse easily, 91
harvesting is not always effective.
Moreover, if spores have already been
produced it will be necessary to repeat
the treatment for several consecutive
years. Chemical control with glyphosate
(in a formulation suited to moist
habitats) is considered effective but
requires regular monitoring. Diquat was
recommended in the past but its use has
been stopped by the European Union in
2002 due to its negative effect on river
ecosystems. A biological control method
has been developed using a beetle species
(Stenopelmus rufinasus) from Argentina
and proved very effective, notably in
South Africa. These beetles were first
released in South Africa in late 1997.
Introduced Range: The red water fern
invades in Europe in the Czech republic,
Belgium, Germany, The Netherlands,
Ireland, Slovakia and in Spain. It also
invades in South Africa, South East Asia,
Australia and in New Zealand.
 Carpobrotus edulis (L.) N.E. Br.
Hottentot Fig, Iceplant
92
Family: Aizoaceae

Basic Description: Carpobrotus edulis is ornamental plant. Today it is planted in

a succulent perennial creeping plant. Its gardens and on sandstone cliffs along
stems are woody and branched and up to the coast. It is not clear when exactly the
2 m long. Roots that penetrate the ground plant was introduced into Israel.
form at each node, so that the plant
Biological and Ecological Properties:
spreads forming dense mats of stems on
Carpobrotus edulis grows in well
the soil surface. Its leaves are opposite,
drained soils, particularly sandy soils
succulent and triangular in cross section.
and sandstone cliffs along the coast.
Leaf color is dark green with reddish
It is sensitive to cold and does not
hues, a trait affected by temperature
tolerate temperatures below -40C. The
and air humidity. Carpobrotus edulis
plant colonizes disturbed habitats, but
flowers are large with a diameter of
can also penetrate undisturbed sites,
6-9 cm. They are yellow, gradually
particularly sand dunes. When it becomes
changing to pink as they age. The fruit
established it grows rapidly at a rate of 1
is succulent and elongated, 2.5-3 cm
m a year. Carpobrotus edulis reproduces
long. It is green and gradually changes
vegetatively and by seeds. Its fruit
to reddish-purple. Seeds are black, about
contains between 1000 and 1800 small
1 mm long. Carpobrotus edulis differs
seeds that are dispersed by opportunistic
from Carpobrotus acinaciformis that has
small mammals, including rats. The seeds
characteristic magenta flowers. The latter
can remain viable in the soil for at least
species is planted in gardens in Israel,
two years. Recent observations in the
but is very rare in natural areas, unlike
field confirmed that in Israel, the plant
Carpobrotus edulis. The hottentot fig
reproduces also by seeds and not only
blooms in Israel in April and May, and
asexually. In the western Mediterranean
occasionally in early summer.
Basin birds have been found to spread
Introduction to Israel: Carpobrotus plant fragments that are used in nest
edulis was introduced into Israel as an building.

Distribution in Israel: Carpobrotus
edulis is invasive along Israel’s coast.
Its populations have become established
in natural areas, both disturbed and
undisturbed. Populations of Carpobrotus
edulis have been found in the coastal
sands along the Mediterranean sea shore
in various protected areas such as the
Na’aman River Mouth Reserve (Galilee
coastal plain), in the Ma’agan Mikhael
Coast and Gador Coast reserves (Carmel
coastal plain), in the Iris Reserve, in the
Poleg Stream Reserve and the Sharon
Coast National Park (northern Sharon).
The numerous populations of Carprobotus
edulis that have been discovered in
recent years along the coast suggest that
the plant may become one of the most
problematic invasive species in Israel’s
coastal ecosystem. Carpobrotus edulis
has been classified as “invasive” in Israel.
Effect on Native Species: Carpobrotus
edulis forms dense mats about 40 cm
thick. This type of growth displaces
other plant species in the infested area.
Carpobrotus edulis reduces concentrations
Geographic Origin: The natural range of Carpobrotus
edulis is in South Africa, along the coast in the Cape
Province.
� References: 1, 32, 55, 59, 162, 194, 284, 302, 340.
of nitrogen, calcium and the pH of the
soil, increasing its acidity and reducing its
fertility. The chemical changes in the soil
prevent germination of native species.
Moreover Carpobrotus edulis stabilizes
sandy areas and modifies the character
of the entire habitat, to the detriment
of psammophilic animals that require
extensive open areas of moving sands.
Carpobrotus edulis is classified as an
“invasive transformer” plant. Yet, a recent
study carried out in Portugal suggests
that the negative effects of the hottentot
fig on native species might be weaker
than previously thought.
Control: Uprooting Carpobrotus
edulis plants is effective if all the plant
fragments are collected and removed.
Since its leaves are succulent and rich 93
in water, fire is not an effective method.
Spraying glyphosate is effective but must
be monitored as plants that seem to
have been killed can recover after a few
months. Moreover, unfocused spraying
could affect native species. No biological
control method by insects or fungi has
been developed yet. Grazing is not an
option due to the high salinity of the
leaves. Restoring an infested site is a long
complex process because the soil remains
acidic for a number of years after the
plants have been removed.
Introduced Range: Carpobrotus
edulis is one of the most invasive
species in nearly all the countries in the
Mediterranean Basin, particularly Spain,
France, Italy and Greece. The plant is
invasive in California and the Azores
Islands. Additional concentrations exist in
southwestern Australia, the southeastern
United States and in New Zealand.
 Conyza bonariensis (L.) Cronquist
Flax Leaved Fleabane
94
Conyza albida Willd. Ex Spreng.
Fleabane
Family: Compositae (Asteraceae)
Basic Description: Conyza bonariensis
and Conyza albida are erect annual
plants. Conyza albida is 1.5-2 m tall and
Conyza bonariensis only reaches a height
of 40-70 cm. Both species have narrow
linear leaves, covered with soft hairs.
Conyza bonariensis has leaves 6 mm
wide, compared to 7-20 mm in Conyza
albida. Leaves and stems are greyish-
green in both species. Conyza bonariensis
branches out, unlike Conyza albida that
has one central stalk. In both species
the inflorescence is at the top of the
branches and comprises white tubular
flowers. Conyza bonariensis has a larger
head, 7-9 mm wide, while Conyza albida
has a smaller head, 2-5 mm across. In
Israel both species bloom from July to
November.
Introduction to Israel: Conyza
bonariensis was first seen in Israel in
1896 whereas Conyza albida was first
recorded only in 1957. There is no specific
information on how these species were
introduced into Israel.
Biological and Ecological Properties:
Conyza bonariensis and Conyza albida
invade similar habitats – disturbed areas
such as roadsides, gardens and fields,
as well as natural areas, mainly riparian
habitats such as riverbanks. Both species
usually invade sites with coarse textured
soils. They grow very rapidly, particularly
Conyza albida that can reach a height of
1.5 m within several weeks. They produce
large numbers of seeds, although no
quantitative studies have been conducted
as for Conyza canadensis. The seeds
are smaller than 1 mm and dispersed by
winds, most about a few dozen meters
from the parent plant, but a small fraction
to much larger distances that can reach
tens of kilometers. Conyza bonariensis
seeds can remain dormant and viable
for 2-3 years. There is no information on
the dormancy period of Conyza albida.
A Greek study found that Conyza albida
possesses allelopathic properties, but it is
not known if the same is true for Conyza
bonariensis.
Conyza bonariensis
Distribution in Israel: Conyza
bonariensis and Conyza albida are known
as “weeds”, particularly the former. That is
to say, they cause damage to agriculture
– Conyza bonariensis in agricultural areas
throughout Israel, and Conyza albida
in the Mediterranean region, south to
Be’er Sheva. The proximity of cultivated
areas to natural areas in Israel has led
to invasion of natural and protected
areas by both species, particularly near
moist habitats. Conyza bonariensis is
invasive along the Alexander and Lakhish
riverbanks, two streams surrounded
by agricultural areas. Both species are
invasive in the Ein Ta’o Nature Reserve
in the Hula Valley, the Ein Yehuda Nature
Conyza bonariensis
Conyza albida
Reserve in the Jordan Valley and even
in the Perat River Nature Reserve in the
Judean Desert. Both species have been 95
classified “invasive” in Israel.
Effect on Native Species: The main
effect of invasive Conyza species is
in displacing native species, mainly
herbaceous species that grow towards the
end of spring. They create shade in dense
stands as well as competing with native
species for soil water. A study has shown
that the presence of Conyza bonariensis
in young fruit tree groves delayed tree
development due to competition for water.
The effect of the allelopathic properties of
Conyza albida on native species has not
been studied yet.
96

Conyza albida

Conyza albida
 97

Conyza bonariensis

Control: The most effective treatment

against both species is manual removal.
The plants, including mature specimens,
are easy to uproot. Mowing encourages
renewal and branching, and reduces
the effect of chemical control methods
applied subsequently. In the early 2000s
Conyza bonariensis developed herbicide
resistance, including to glyphosate, the
most common agricultural herbicide
used against invasive species.
Nevertheless, chemical control is still
considered effective if the herbicide
is sprayed in the early stages of plant
development, at the rosette stage, before
the central stem develops. No means of
biological control has been developed
against either of these species. It has
been observed, particularly in the Perat
River Reserve, that Conyza bonariensis
and Conyza albida tend to dominate sites
from which other invasive plants, such
as Pennisetum clandestinum, have been
removed.
Introduced Range: Conyza bonariensis
is invasive in California, China, Central
Europe, the Mediterranean Basin, South
Africa and Australia. Conyza albida is
invasive in all of Australia, Southeast
Asia and in most of the Pacific Ocean
islands.

Geographic Origin: Both Conyza species originate

in tropical South America. Since they are invasive
throughout the world, particularly in the American
continent, it is difficult to precisely define their natural
range.

� References: 61, 63, 88, 173, 249, 299, 302, 304.

 Conyza canadensis (L.) Cronquist
Canada Horseweed
98
Family: Compositae (Asteraceae)
Basic Description: Conyza canadensis is
an erect annual plant, 1-2 m tall. In most
cases it has a lone, unbranched central
stem. Leaves are narrow and linear, no
wider than 6 mm. Its leaves and stem are
light green. Unlike other invasive Conyza
species in Israel, Conyza canadensis
leaves are almost completely hairless.
The upper part of the plant bears dozens
of inflorescences at the head of the plant,
each with 50 whitish-pink ray flowers.
Flower head size is 2-3 mm. In Israel
Conyza canadensis blooms from July to
November.
Introduction to in Israel: Conyza
canadensis was first found in Israel in
1939 in the Upper Galilee. Its seeds may
have been unintentionally introduced in
special food for fish or it may have been
dispersed in the air from neighboring
countries.
Biological and Ecological Properties:
Conyza canadensis invades disturbed
areas such as roadsides, gardens and
cultivated fields, but also in natural
sites such as stream banks and around
marshes. It prefers coarse-textured soils
with a pH of 4-10. Conyza canadensis can
grow in soils with a high salinity level.
It grows rapidly and plants can grow to
over 1.5 m within a number of weeks. It
produces a very large number of seeds:
research has found that one individual can
produce 130-200,000 seeds or more. The
seeds are very small and dispersed by
wind. About 99% of Conyza canadensis
seeds are dispersed up to about 900
m from the parent plant. Some of the
seeds fly up to heights of several dozen
meters and can distance themselves up
to 500 km from the parent plant. The
seeds remain viable for a relatively short
time, usually 2 to three years, although
one study found Conyza canadensis seeds
that were viable 20 years after dispersal.
Studies have shown that the seeds do
not germinate if placed deeper than 0.5
cm below the soil surface. A recent study
conducted in Serbia showed that the plant
releases phenolic compounds in the soil
that have a long-lasting negative effect on
seed germination and seedling growth of
native herbaceous species.

Distribution in Israel: Conyza
canadensis is classified as a weed in
cultivated fields in Israel’s Mediterranean
region, south down to Be’er Sheva. It
also creates dense stands after herbicide
spraying in JNF planted pine forests.
As a result of the proximity between
agricultural areas and planted forests, all
alien Conyza species, including Conyza
canadensis, have invaded natural and
protected areas, particularly in riparian
habitats. For example Conyza canadensis
has spread along the stream banks of
the Alexander River and the Lakhish
River that are surrounded by agricultural
areas. It also invades the Ein Ta’o Nature
Reserve in the Hula Valley, the Ein Yehuda
Nature Reserve in the northern Jordan
Valley and even the Perat River Reserve
in the Judean Desert. Conyza canadensis
has been classified “invasive” in Israel.
Effect on Native Species: The
main effect of Conyza canadensis
Geographic Origin: The natural range of Conyza
canadensis includes most of the United States and
Canada, except for Canada’s northern region.
� References: 61, 63, 67, 68, 75, 145, 173, 227, 249, 297, 299, 302, 304, 339, 360.
concentrations is in displacing native
species, mainly herbaceous species that
grow towards the end of spring. The
density of infested sites may be as high
as 130 plants per sq. m., which creates
shade and suppresses the development
of native species. Conyza canadensis
also uses soil water at the expense of
other species. Research has shown that
the presence of Conyza canadensis in
fruit tree groves delayed development in
young trees due to competition for water
so it presumably impacts native species
the same way. The strong allelopathic
properties of the plant reduce abundance
and cover of native species in infested
areas.
Control: The most effective treatment
against Conyza canadensis is manual 99
removal. The plants, including mature
specimens, are easy to uproot. Mowing
encourages renewal and branching,
and makes subsequent application of
chemical control methods difficult. In the
late 1990s Conyza canadensis apparently
developed resistance to herbicides,
including to glyphosate. Nevertheless,
chemical control is still considered
effective if the herbicide is sprayed in the
early stages of plant development, at the
rosette stage, before the central stem
develops. No means of biological control
has been developed yet, although there
are a number of studies underway.
Introduced Range: Conyza canadensis
is invasive throughout Europe, in
temperate Asia and in the Mediterranean
Basin.
 Cyperus involucratus Rottb.
Umbrella sedge
100
Family: Cyperaceae
Basic Description: Cyperus involucratus
is a perennial herb 50-180 cm tall,
completely smooth, with many thick erect
ridged stems. The leaves are light brown
sheaths 10-20 cm long. The stems bear
umbelliform inflorescences, composed of
14-24 straight and narrow leafy bracts,
10-30 cm long and 4-17 mm wide. The
bracts around the inflorescence are
rough, with protruding veins, spread
horizontally and drooping slightly at the
edges, giving the plant an umbrella-
like appearance. The spike bears 6-18
compressed spikelets, ovoid or oblong,
3-12 mm long and 1.5-3 mm wide. Each
spike bears 8-36 tiny yellow-greenish or
light brown flowers 2 mm long and 1.1
mm wide. In Israel the species blooms in
summer. Cyperus alternifolius is a similar
species that can be differentiated from
Cyperus involucratus by its longer spikes
(5-20 mm).
Introduction to Israel: Cyperus
involucratus was introduced into Israel
as an ornamental and planted in many
gardens. The exact date of introduction is
unknown.
Biological and Ecological Properties:
There is almost no information relating
to the ecological traits of Cyperus
involucratus. The species grows in moist
habitats such as riverbanks and marshes
and apparently reproduces mainly
asexually. Pieces of rhizomes that break
off the parent plant are carried by flowing
water and root in new sites creating new
plants downriver, a means of dispersal that
enables Cyperus involucratus to spread
rapidly. According to one source the species
is considered resistant to frost, hot dry
weather and even salinity. It is not known
to have any specific soil requirements.
Its natural range and the areas it has
invaded testify to its resistance to high
temperatures. Cyperus involucratus does
not require particularly disturbed areas to
become established, and can even do so in
the midst of dense native vegetation, an
indication of its great competitiveness.
Distribution in Israel: Cyperus
involucratus invades along the banks of the
Perat River in the northern Judean Desert.
It also grows at the Yarkon River sources
and was reported in 2011 along the
banks of the Sorek River in the Jerusalem
Mountains. The earliest report of Cyperus
involucratus in natural areas in Israel are
from 2004 in the Jerusalem Mountains.
There have been reports of specimens in
the area of Lake Kinneret but these have
not been verified and may probably be
specimens of Cyperus alternifolius, which
is very similar to Cyperus involucratus.
The rapidity of its invasion is apparent
along river courses: during seven months
between July 2009 and January 2010,
dozens of new specimens appeared
along the banks of the Perat River. Since
this sedge species is widely used as an
ornamental in Israel it will very probably
invade new sites in the country in the near
future. At this point Cyperus involucratus
is still considered a “naturalized” species in
Israel since there are no records older than
10 years, but it will certainly be regarded
as “invasive” in the next few years.
Effect on Native Species: Cyperus
involucratus forms extremely dense stands
that displace native plants. The numerous
stands of this species can dramatically
Geographic Origin: The natural range of Cyperus
involucratus is in East Africa.
� References: 47, 100, 139, 162, 330.
modify the character of riverbank
vegetation in the areas it invades.
Control: There is no information on 101
control methods for Cyperus involucratus.
A preliminary experiment in the Perat
River Nature Reserve indicates that young
plants can be easily uprooted, while they
still have shallow root systems. Larger
clusters of the plant were successfully
uprooted with the help of a backhoe
loader. However some foci may not be
accessible to heavy machines. Its invasion
in wetlands and along rivers complicates
the possibility of spraying herbicides.
Future studies will have to examine what
management methods are the best suited
to control this alien sedge species.
Introduced Range: Cyperus
involucratus is invasive in the southern
United States, particularly in California,
Arizona, Texas, Louisiana, Mississippi
and Florida. It is also invasive in Mexico
and the Caribbean Islands. Cyperus
involucratus is considered “invasive” in
Hawaii and most of the Pacific islands.
It is also invasive in West Australia and
northern New Zealand.
 Cyperus odoratus L.
Fragrant Flatsedge
102
Family: Cyperaceae

Basic Description: Cyperus odoratus its seeds were brought in inadvertently in

is an annual, sometimes perennial feed mixes for birds.
grass, 10-50 cm tall, with a smooth,
Biological and Ecological Properties:
erect stem, 1-4 mm, triangular in cross-
There is very little information on the
section. The rhizome is short, 0.5-2 cm
ecological features of Cyperus odoratus.
long and only 0.5-1.5 cm wide. At the
In its natural range the species grows
base of the plant are 3-6 linear leaves,
in dense clumps in moist habitats,
10-65 cm long and 4-12 mm wide. The
particularly along coasts and on
leaves are canaliculate and have slightly
riverbanks. Cyperus odoratus reproduces
rough edges. The sheaths are green or
by seeds only. The seeds float on the
purplish. The stems bear umbelliform
water surface and are thus dispersed to
inflorescences composed of 5-8 linear
considerable distances. It can penetrate
leafy bracts that are spread horizontally
and establish itself in dense native
or at a 30-600 upwards angle. The
vegetation, even in undisturbed areas.
peduncle is 2-8 cm long and bears 1-
5 spikelets 10-20 mm long and 10-30 Distribution in Israel: Cyperus odoratus
mm wide. In Israel the species blooms was first reported in 2006 in the area of
throughout the summer until October. Lake Agmon in the Hula Valley, on the
water’s edge in flooded areas. In 2008
Introduction to Israel: The first official
it was reported in the coastal sands in
record of Cyperus odoratus is from 2006.
the southern coastal plain. In 2009 it
According to one source Cyperus odoratus
was reported in the same type of habitat
was already found in 1980 but was not
in the northern Sharon region. Cyperus
documented. The source of introduction
odoratus is considered a “naturalized”
into Israel is unclear. The species is
species in Israel.
a pond ornamental in North America and
may have been imported into Israel for Effect on Native Species: Cyperus
this purpose. Another possibility is that odoratus forms dense stands that displace
 103

native species, particularly those that define the optimal methods of control and
grow on the banks near the waterline. management of Cyperus odoratus.

Control: There is no information on Introduced Range: Cyperus odoratus is

methods of controlling Cyperus odoratus. considered an invasive species in Europe.
Since it has a small shallow root system, In the tropics it is known as a troublesome
uprooting should be possible and species in roadside ditches, agricultural
effective. Future studies will have to areas and disturbed moist habitats.

Geographic Origin: Cyperus odoratus has a pan-tropic

distribution. It grows in tropical regions of Asia and
Africa, including Madagascar, Australia, the Pacific and the
Americas. In the New World it is also common beyond the
tropical region, from California to southeast Canada.

� References: 62, 66, 85, 330.

 Datura stramonium L.
Common thornapple
104
Family: Solanaceae
Basic Description: Datura stramonium
is a large, branching annual, 50-130 cm
tall. Its leaves are green, sometimes light
purple, glabrous or with sparse hairs.
Leaves are simple, glabrous, alternate,
oblong or triangular. The leaf blade
is irregularly lobed and its margin is
unevenly serrated. The leaves are 5-20
cm long, sometimes reaching 35 cm and
4-15 cm wide. The flowers are white or
occasionally a bit purple, large and funnel
shaped. They are up to 20 cm long and
have 5 fused petals each of which ends in a
long point. The flowers close at night. The
fruit is a rounded elongated capsule, with
a diameter of 2-5 cm, usually covered in
short (about 15 mm) spines. The capsule
opens into four sections and releases
the seeds that are black or dark brown,
kidney shaped, 3-3.5 mm long. The
capsule can hold 600-700 seeds. In Israel
Datura stramonium blooms from June
through October.
Introduction to Israel: Datura
stramonium was first recorded in Israel in
1912. There is no information on how it
was introduced into Israel. Its leaves were
used in traditional medicine in the East to
treat malaria, and this may be why it was
imported into Israel.
Biological and Ecological Properties:
The preferred habitat of Datura
stramonium is riverbanks and other moist
habitats. The species has a very well
developed horizontal and vertical root
system. It reproduces only by seed. Non-
dormant seeds will germinate three days
after being watered. A mere 10 mm of
rain is sufficient for Datura stramonium
seeds to germinate. Growth in early
stages is very rapid and the plant can
grow up to 2 cm daily, and flower a mere
3-5 weeks following germination. There
are no studies on which insects pollinate
Datura stramonium plants growing in
Israel, but in South Africa pollination is
carried out mainly by moths and bees. An
individual plant can produce up to 30,000
seeds, which may remain dormant and
viable for 40 years or more. The plant has
allelopathic properties.
Distribution in Israel: Datura
stramonium invades along rivers in the
Mediterranean and arid regions, in sites
such as the Sorek River Nature Reserve
in the Jerusalem Mountains, at the mouth
of Wadi Kidron in the Dead Sea Valley,
in the lower section of Wadi Og and in
the Perat River Nature Reserve. Datura
stramonium is also invasive in the Hula
Nature Reserve in the Hula Valley. It
grows in irrigated agricultural areas and
on roadsides in Israel’s Mediterranean
region. Datura stramonium is classified as
“invasive” in Israel.
Effect on Native Species: The large
leaves of Datura stramonium create
extensive shade that displaces native
species. It can form dense stands locally
along riverbanks, in which native plant
species are absent. Datura stramonium
also possesses allelopathic properties:
it releases chemical compounds from its
seed coat that prevent germination and
growth of other plant species.
Control: It is possible and advisable to
105
uproot young Datura stramonium plants
before they produce seeds. Uprooting
adult plants is more difficult. Chemical
treatment is only effective against young
specimens. Herbicides such as 2,4-D or
glyphosate are considered effective, but
spraying may prove problematic since
most of the species grow near the water.
There is no effective biological control
method against Datura stramonium.
Introduced Range: Datura stramonium
is invasive in many parts of the world
including temperate northern Europe,
northern Canada and even Alaska. There
are large invasive sites in South Australia
and the southern and eastern parts of
South Africa and the Galapagos Islands.

Geographic Origin: The natural range of Datura

stramonium has not been defined precisely. Apparently
it originates in Central America or tropical South
America.

� References: 61, 85, 104, 149, 162, 239, 283, 334, 340.
 Dodonaea viscosa L.(Jacq.)
Hopbush, Varnish Tree
106
Family: Sapindaceae

Basic Description: Evergreen shrub, of the specimens that invade natural

1-3 m tall with simple, relatively large
elongate leaves, 5-10 cm long and
0.8-1.2 cm wide. The leaves have
conspicuous veins and their edges may
be weakly toothed or undulating. The leaf
tip is round or pointed; leaves secrete
a gummy exudate which gives them
a shiny appearance and makes them
sticky. Dodonaea viscosa is dioecious, i.e.
it has separate male and female plants,
or polygamodioecious, i.e. male and
bisexual on one individual, and female
and bisexual on another. The flowers are
inconspicuous, (less than 1 cm across),
and lack petals. The flowers are pale
green, the sepals are greenish-yellow
and stamens are brown. Female flowers
are wind pollinated. After pollination the
individuals recorded in natural habitats
in Israel develop into 2 to 3 winged
papery capsules; wings are pale brown
or coral pink and capsules turn red or
purple as the fruit matures. Each capsule
encloses 2 to 3 smooth black seeds. The
Dodonaea viscosa complex includes up to
7 subspecies. The morphologic properties
habitats in Israel are characteristic of
the subspecies viscosa, the only member
of the complex with distinctive 2- and
occasionally 3-winged capsules.
Introduction to Israel: There is no
reliable data relating to the time when
Dodonaea viscosa was introduced
to Israel. It is possible that the first
individuals were introduced for their
medicinal properties. The shrub was used
in the Mediterranean Basin for hedges
and it may have been introduced for this
purpose as well. It is also considered an
attractive garden plant. Sterile hybrids of
Dodonaea viscosa and other Dodonaea
species were cultivated in Israel and
elsewhere and are today favored
waterwise ornamentals.
Biological and Ecological Properties:
Dodonaea viscosa is an extremely hardy
species, tolerant of salinity, drought
and pollution. It can grow on a variety
of soils, including on sand dunes in
coastal environments. It grows fast and
is tolerant of strong winds. The species
produces large quantities of seeds, yet
no study has quantified this aspect.
Seeds are dispersed by wind. There
are no observations related to fruit or
seed predation by native organisms
in Israel. Reproduction in Israel is
apparently only by seeds. According to
our observations in Israel the plant is
able to establish itself in undisturbed
habitats such as rocky slopes in the
mountainous area, and on coastal
sands where it forms dense thickets.
Dodonaea viscosa shrubs are able to
resprout from the base.
Distribution in Israel: Isolated
individuals have been recorded along
the banks of the Kelah Stream on the
northern part of Mount Carmel and in
the area of Kibbutz Bet Oren located in
the same area. The presence of these
individuals is not recent and may be
a relict of former cultivation when the
shrub was grown as a hedging plant.
In the past 5 years or so new foci
of Dodonaea viscosa were reported,
mainly on coastal sands, for example
in the Rishon LeTsiyon sands, south
of Tel Aviv as well as on calcareous
sandstone (kurkar) hills near Nes
Tsiyona in the same region. Surprisingly
the plant created dense stands with tall
individuals that became established in
undisturbed sites. Several individuals
were found in 2011 in the Ramat
Hanadiv protected park located on
the southern tip of the Carmel ridge;
however these individuals are still
young seedlings and their number is
restricted. Dodonaea viscosa has been
classified as “invasive” in Israel.
Effect on Native Species: Dodonaea
viscosa forms dense thickets in the
coastal sand dunes that displace all
native plants such as Retama raetam
or Artemisia monosperma. The plant’s
dense foliage creates a thick litter of
dead leaves beneath mature individuals
and the understory completely lacks
other plant species. The fact new foci
recently emerged in different locations
in undisturbed habitats is a reason for
concern, particularly for coastal sandy
habitats.
Control: There is virtually no information
about control methods that may be
efficient against Dodonaea viscosa.
107
Physical or mechanical treatments
are probably useless as the plant
readily resprouts after cutting. Further
experiments will be necessary in order to
evaluate the plant’s reaction to targeted
chemical control. There is no method of
biological control.
Introduced Range: Dodonaea viscosa
has been introduced on all continents,
except Antarctica, and its distribution
extends from 440S in the South Island of
New Zealand, to 330N in California and
Arizona. It is planted as a decorative
shrub in South Africa, Australia and
southern U.S.A. but the plant was not
reported invasive in these countries. The
only country where Dodonaea viscosa is
officially considered invasive, apart from
Israel, is Cyprus.

Geographic Origin: The precise natural range of

Dodonaea viscosa is uncertain. Dodonaea viscosa is
probably native to Australia, although some sources
consider it is native also in India and tropical Africa. The
subspecies viscosa is found only as a mangrove associate
or on sand dunes close to the foreshore, along the
northeastern Queensland coast and offshore islands.
� References: 28, 142, 148, 176, 342
 Eichhornia crassipes (Mart.) Solms
Water Hyacinth
108
Family: Pontederiaceae
Basic Description: Eichhornia crassipes
is a floating aquatic plant with glabrous
kidney-shaped, dark green shiny leaves,
15 cm long and 10 cm across. The petiole
is swollen and its diameter is 2-3 cm and
has a spongy consistency that allows the
leaves to float on the surface of the water.
Its erect inflorescence is about 30 cm
long and has 5-8 large purple flowers. The
base of the upper petal is light blue and
it has a yellow spot on it. Root diameter
is 1-2.5 cm and roots can reach a length
of 1 m. In Israel it blooms from early July
through mid-September.
Introduction to Israel: Eichhornia
crassipes was introduced into Israel as an
ornamental plant, although it is not clear
when.
Biological and Ecological Properties:
Eichhornia crassipes possesses a
combination of many of the ecological
characters of invasive plants: it
reproduces from seeds, but mainly
vegetatively by stolons. The underwater
stems develop leaves that in time break
off from the parent plant and become
independent plants. The reproductive rate
of this species is extremely rapid: the
number of plants in a given population
can double itself every 5 days. One single
plant produces enough biomass to cover
an area of 600 sq.m. The considerable
amount of vegetative reproduction
creates “rafts” that move along the
water surface with the current creating
new invasive sites along shores and on
riverbanks. New specimens flower within
3-4 weeks of their formation. Its fruit can
contain up to 300 seeds. A recent study
in South Africa showed soil seed density
may reach as much as 2500 seed/m2.
The seeds sink, germinate and develop
roots, detach themselves from their roots
and float on the water surface. They can
lay dormant in the dry mud for up to 15
years. Eichhornia crassipes seeds stick to
the feet of water birds, which probably
play a significant role in their dispersal.
Nevertheless, humans are their principal
dispersal vector, e.g when emptying
infested aquarium contents into water
bodies or when translocating plant parts
from one infested water body to another
area. In sub-tropical areas, such as
Israel, freezing winter temperatures harm
Eichhornia crassipes leaves, but do not
affect submerged parts from which the
plant regenerates in spring.
Distribution in Israel: The first record
of the species in Israel is from 1953-1954
when it invaded and completely choked
the Yarkon River in the southern Sharon
near Tel-Aviv. A second invasion occurred
in 1961-62 at the mouth of the Alexander
River in the northern Sharon. The foci
reported in the Yarkon River in the 1950s
and in the early ‘60s remained isolated
events and did not last. Then, in the early
years of 2000 the water hyacinth began
to invade again, this time in several sites
throughout the country. In late 2003 a
large population that completely covered
a water pool extending over 3.9 ha in
an abandoned sandstone quarry in the
northern Sharon region was reported. Low
winter temperatures in 2007-2008 did
not affect this population which remains
stable up to now. During 2005-2006 the
plant began to invade the Alexander River
in the northern Sharon and for a while
covered a small patch on the river. Winter
floods swept the plants into the sea, but
in 2009 the species once again returned
to the river. In 2006 another population
was found in the Na’aman River in the
Galilee coastal plain, and despite manual
removal the plant regularly spreads again
in this stream. In 2005 a small focus
was reported in the Gedora River in the
same region and since 2009 the plant
chokes extensive sections of the river.
There is a stable 8-year old population in 109
a drainage channel near the Hula Nature
Reserve in the Hula Valley. In September
2011 a small population was found in the
Arbel Stream in the Arbel Nature Reserve
located in the eastern lower Galilee.
In view of the stability of populations
in slow-flowing water bodies, and the
multiplication of invasive sites in various
areas during the last decade, Eichhornia
crassipes has been classified as “invasive”
in Israel. There is serious danger that
the plant will invade Lake Kinneret in the
future. The water hyacinth is still sold as
an ornamental in plant nurseries in Israel.
Effect on Native Species: When
Eichhornia crassipes covers a water body
the abiotic properties of the infested
ecosystem change drastically: the pH
decreases, dissolved oxygen and the
amount of light in the water are reduced.
These changes eventually eliminate
native water plants, fishes and other
aquatic organisms. As a result of the
high transpiration rate of Eichhornia
crassipes, the rate of water evaporation
from the surface of a water body covered
by this species is eight times the amount
evaporated from a non-infested water
body. Consequently the proliferation
of Eichhornia crassipes can lead to the
total collapse of a freshwater ecosystem,
110
 111

particularly in sluggish water bodies. insects. A recent study recommends

Considering the nature and the extent of the
damages it causes to infested ecosystems
Eichhornia crassipes is considered an
“invasive transformer”.
Control: The simplest and most suitable
control method for small foci is manual or
mechanical collection. Chemical control with
glyphosate is feasible only with formulations
adapted to aquatic environments. A variety
of biological control methods using insects
has been developed, of which the most
effective are two beetle species Neochetina
eichhorniae and Neochetina bruchi.
However, biological control in subtropical
regions is not very effective due to the
low winter temperatures that affect the
combining chemical control (glyphosate)
with biological methods. Another method
tested lately is biological control using
chemicals found in the leaves of Lantana
camara, another invasive species.
Introduced Range: Eichhornia crassipes
is one of the 100 most invasive organisms
in the world according to the IUCN’s
Invasive Species Specialist Group (ISSG).
The species invades mainly tropical and
subtropical regions, but also temperate
areas such as Australia, the southern
and western United States, South Africa,
tropical Africa (Lake Victoria), India,
Southeast Asia, southern Europe, New
Zealand and Canada.

Geographic Origin: The natural range of Eichhornia

crassipes is the Amazon basin in northern Brazil.

� References: 6, 9, 32, 85, 169, 178, 236, 239, 340, 349, 350.
 Eucalyptus camaldulensis Dehn.
112
Murray Red Gum, River Redgum
Family: Myrtacea
Basic Description: Eucalyptus
camaldulensis is a broad-leaved,
evergreen tree species, whose height is
10-35 m. Its bark is whitish-grey and
is shed in strips. Its leaves are multi-
shaped, but usually oblong, lanceolate
with pointed tips and smooth edges. The
leaf is glabrous, hard and both sides are
identical. Its greenish-greyish leaves
are 7-9 cm long and 1-1.5 cm wide.
The flowers (1-1.5 cm) lack petals and
have many cream-colored stamens. The
inflorescence is a short panicle with 7-11
flowers. The fruit is a conical brown pod,
6-7 mm long. Upon ripening it opens
and releases tiny seeds. Eucalyptus
camaldulensis blooms twice yearly in
Israel – from March to May and from
September until early November.
Introduction to Israel: Jewish settlers
introduced Eucalyptus camaldulensis into
Israel around 1890 to drain swamps. The
first trees were planted in the area of
Hadera in the northern Sharon, Rosh Pina
in the Hula valley and Petah Tikva in the
southern Sharon. During the 1920s the
Jewish National Fund and British Mandate
officials planted the species in many parts
of the country. Eucalyptus camaldulensis
is the most common planted species in
Israel after Pinus halepensis and Pinus
brutia.
Biological and Ecological Properties:
The natural range of Eucalyptus
camaldulensis in Australia gives an
indication of this species great ecological
plasticity (see below ‘Geographic Origin’).
From plantings in Israel we can see that
Eucalyptus camaldulensis can grow in
saturated soils in the Mediterranean
region, in dry habitats in semi-arid
areas, in heavy clay soils and even in
sandy soils. The species is resistant to
moderate soil salinity levels. It has a well
developed deep root system, reproduces
exclusively from seeds and is pollinated
by bees. It grows very rapidly in moist
habitats, reaching heights of about 6-
8 m within a few years. Eucalyptus
camaldulensis can usually produce seeds
at the age of 5-7 years, and sometimes
even earlier. In its natural range the
species can live between 500 and 1000
years. It regenerates after cutting but is
 Tavor River reserves in the eastern lower
Galilee and in the Tel Dan Reserve in the
Hula Valley. Eucalyptus camaldulensis has
been classified as “invasive” in Israel.

Effect on Native Species: The effect

sensitive to fire. Eucalyptus camaldulensis
has allelopathic properties: a number of
studies have shown that its fallen leaves
release chemical compounds that prevent
germination of other plant species.
Distribution in Israel: Until recently
Eucalyptus camaldulensis was thought
not to reproduce from seeds in Israel,
and that all existing specimens had
been planted. But it turns out that many
of the trees growing in moist habitats
developed from seeds and formed new
populations. Eucalyptus camaldulensis
invades natural areas, some protected,
such as the Kadima Sands National Park
in the northern Sharon, the Acacia albida
Reserve near Ashdod in the southern
coastal plain, the Judean Mountains
National Park (on the banks of the
Kesalon Stream), and the Sorek River
Reserve in the Jerusalem mountains,
the Kare Na’aman Reserve in the Galilee
coastal plain, the Yavne’el River and the
of Eucalyptus camaldulensis on native
species in Israel has not yet been
investigated. However local surveys in large
concentrations of Eucalyptus camaldulensis
show a very low number of native species,
both herbaceous and ligneous. The scarcity
of local species in the understory of
Eucalyptus camaldulensis is presumably due
to its allelopathic properties. In addition,
establishment of the invasive tree in habitats
with high water tables is likely to lower the
groundwater level and may eventually lead
to partial drainage of the infested habitat.
Future studies will have to examine these 113
aspects.
Control: Small seedlings and saplings
can be uprooted. Cutting alone is not
effective because Eucalyptus camaldulensis
regenerates from stumps. Chemical control
by cut-stump or drill-fill with triclopyr
kills mature trees. Since Eucalyptus
camaldulensis is one of the most commonly
planted tree in the world, no biological
control methods have been developed
against it for fear of harming eucalyptus
plantations.
Introduced Range: Eucalyptus
camaldulensis is invasive in most parts of
South Africa particularly in the Western
Cape Province where it has been designated
invasive and may not be planted near water
sources. Eucalyptus camaldulensis is also
invasive in California and Hawaii.

Geographic Origin: Eucalyptus camaldulensis is

endemic to Australia. It grows almost throughout the
continent, with the exception of the western section
of South Australia and the southeastern section of
Western Australia. Its natural range extends over an
extremely varied range of climatic regions, from the
humid-tropic region in the north to the arid area of the
central continent and down to the Mediterranean areas
in southeastern and southwestern Australia.
� References: 4, 40, 58, 85, 149, 190, 267, 300, 302.
 Ficus benghalensis L.
East Indian Figtree, Indian Banyan
114
Family: Moraceae
Basic Description: Ficus benghalensis
is an evergreen monoecious tree, 15-27 m
tall, with support roots growing from
its branches. Its dark green leaves are
ovate and oblong, 10-30 cm long and
7-20 cm wide. The leaf tip is blunt and its
margin entire. The leaves are hard and
leathery, with a glabrous upper side and
a hairy underside. The petiole is hairy and
1.5-7 cm long with a diameter of 5 mm.
The flowers are minute, monosexual and
arranged in paired rounded hairy red
syconiums in the leaf axils. The syconium
has a diameter of 1.5-2 cm.
Introduction to Israel: Ficus
benghalensis was introduced into Israel
as an ornamental, presumably in the
1940s, but the exact introduction date
remains unknown. Due to its tolerance for
high temperatures it has been planted in
gardens in arid areas of the country, such
as the Dead Sea Valley.
Biological and Ecological Properties:
Ficus benghalensis is tolerant of high
temperatures and is apparently sensitive
to low temperatures. In the areas it
invades in Israel it reproduces by seeds
that germinate and become established in
rock fissures. Pollination is by a tiny wasp
(Eupristina masoni) that can penetrate
the syconium. Seed dispersal is mainly
by birds. One of the main dispersal
agents seems to be the Tristram’s grackle
(Onychognathus tristramii), a native
bird species that can disperse the seeds
to at least 3.5 km from the parent tree.
Seeds have also been found in porcupine
(Hystrix indica) droppings in the Dead
Sea area indicating that this rodent is
also involved in Ficus benghalensis seed
dispersal. The tree can renew itself from
its roots as can be seen in trees damaged
by floods in desert seasonal streams.
Distribution in Israel: Ficus
benghalensis geminates and grows in
moist habitats in seasonal stream beds
and cliffs in the arid areas in the Dead
Sea Valley. It is invasive only in the
Dead Sea area in Israel, mainly along
the Arugot and David rivers in the Ein
Gedi Nature Reserve and along Wadi
Bokek in the southern Dead Sea Valley.
The first seedlings were found in 1996
and since then several dozen mature
specimens have become established.
A survey conducted in 2001 found ca.
500 individuals, mostly seedlings and
saplings, along the Arugot River alone.
Ficus benghalensis is the most common
of the three alien Ficus species that
invade the area. Its invasion rate seems
to be extremely rapid and requires
urgent management and control before
the proliferating individuals start to set
seeds. Ficus benghalensis is classified as
“invasive” in Israel.
Effect on Native Species: Ficus
benghalensis can displace native species,
particularly those growing near stream
beds, thanks to its broad canopy and
large leaves that create shade. A number
of animal species feed off the tree but
there is still no information on its effect
on the native fauna in Israel.
Control: Mechanical control could be
effective against Ficus benghalensis
seedlings only if the entire root system
Geographic Origin: The natural range of Ficus
benghalensis, according to most experts, is in India’s
arid region where it grows in low places, at altitudes up
to 600 m.
� References: 29, 30, 100, 261, 315.
that penetrates rock crevices can
be removed. Eradicating established
individuals requires focused chemical
control with the herbicide triclopyr that 115
is very effective against Ficus trees. This
treatment must be applied with great
care due to the proximity to water. Seeds
sometimes germinate in sites such as cliff
crevices that are difficult to reach which
complicates treatment even more. There
is no biological control method against
this species. In order to reduce the
number of seeds in the region and as part
of the management program it will be
necessary to uproot all existing individuals
in the Dead Sea area and to forbid further
planting in the region.
Introduced Range: Ficus benghalensis
is invasive in Florida since the 1980s.
In Dade County, Florida, it is forbidden
to plant the species. The tree is also
invasive, albeit on a small scale, in
northeastern and central Queensland in
Australia.
 Ficus microcarpa L.fil.
Chinese Banyan, Curtain Fig
116
Family: Moraceae
Basic Description: Ficus microcarpa is
an evergreen monoecious tree, 15-20
m tall, with thin aerial roots growing from
its branches. Its bark is grey and smooth.
Its dark green shiny leaves are glabrous,
relatively rigid, alternate, elliptical or
oblong, 5-8 cm long (occasionally longer)
and 3-5 cm wide. The petiole is 0.6-2 cm
long. The flowers are minute, monosexual
and arranged in paired rounded syconiums
in the leaf axils. The syconia are smooth,
red, sometimes yellow, and blacken as
they mature. The syconium diameter is a
mere 6-10 mm. The seeds are smaller than
1 mm.
Introduction to Israel: Ficus microcarpa
is an ornamental in Israel and the world
over. Its introduction date is unknown. It
was planted mainly in the coastal region,
usually along streets and boulevards.
Biological and Ecological Properties:
The natural range of Ficus microcarpa
suggests that it requires habitats that
are warm and moist throughout the year.
The tree reproduces by seeds that can
germinate and establish themselves in
rock crevices with no soil pockets. Ficus
microcarpa grows rapidly. Pollination is
by the wasp Parapristina verticillata that
invaded Israel in the late 1980s, or possibly
even earlier. Examination of Tristram’s
grackle (Onychognathus tristramii)
droppings, a native bird species in the Ein
Gedi area, in the Dead Sea Valley, indicate
that this local species plays a major role
in dispersing Ficus microcarpa seeds.
According to evidence from other parts of
the world bats and rodents also disperse
Ficus microcarpa seeds. The tree can
recover rapidly after cutting by renewal
from its roots.
Distribution in Israel: Of the three
invasive Ficus species in Israel, Ficus
microcarpa is the least common. A number
of trees were first discovered in the Ein
Gedi Nature Reserve, in the Dead Sea
Valley, in 1992. Since then a number of
large specimens have become established
in the midst of the dense vegetation near
the waterfalls. Many small individuals grow
along the David River that flows in the Ein
Gedi Reserve. The trees growing on rocks
in the stream bed are damaged by winter
floods, but renew themselves afterwards.
Some of the trees growing near relatively
large falls seem not to be affected by
the floods. The Dead Sea region is the
only area where the tree was reported
invasive in Israel. There is a risk that the
invasion rate will accelerate significantly
when the seedlings and saplings that
have already established themselves will
begin to produce seeds within the natural
protected area. F. microcarpa is classified
as “invasive” in Israel, but only in the Dead
Sea Valley region.
Effect on Native Species: Ficus
microcarpa can form dense stands that
displace native species that grow near
stream beds in oases. It can penetrate and
become established in dense vegetation,
a trait that improves its ability to displace
native plant species.
Control: Uprooting Ficus microcarpa
seedlings is effective if all the roots
penetrating rock crevices are removed.
Often this is not possible and trees
regenerate from their roots. Established
trees should be controlled chemically by
concentrated application of triclopyr, which
is very effective against Ficus species. This
treatment must be applied with great care
due to the proximity to water. It is very 117
difficult to implement chemical control for
trees growing on high cliffs, which can
only be reached with climbing equipment.
No biological control method has been
developed yet for this species. In order to
prevent further invasion in the Deas Sea
Valley it is necessary to uproot all the plants
growing in gardens in the area. They can be
replaced by native tree species that have
equivalent scenic value and shade, but do
not threaten the natural vegetation of the
Ein Gedi Oasis.
Introduced Range: Ficus microcarpa is
invasive in Florida since the 1970s in the
wake of its pollinating wasp. Since then
it has become invasive in Mexico, San
Salvador, Honduras, Nicaragua and Panama.
In the 1980s the species began invading
South America – Colombia, Venezuela,
Bolivia and Ecuador. Ficus microcarpa is
also invasive in a number of Pacific islands,
particularly Hawaii, where it began invading
since the late 1980s.

Geographic Origin: The natural range of Ficus

microcarpa is very broad and extends from India and Sri
Lanka to southern China, Cambodia, Vietnam, Indonesia
and northern Australia. Ficus microcarpa grows in low
altitude rainforests, river banks, flooded flats, marshes
and cliffs in these areas.
� References: 29, 30, 315, 340.
 Ficus religiosa L.
Bo Tree, Sacred Fig
118
Family: Moraceae
Basic Description: Ficus religiosa
is an evergreen or semi-decidous
tree, monoecious, 6-15 m tall that
has no supporting roots growing from
its branches. Its bark is grey and
furrowed. The light green leaves are
ovate-orbiculate, 6-15 cm long and
4-11 cm wide. At the tip of the leaf is
a narrow point, up to 5 cm long, which is
characteristic of Ficus religiosa leafs. The
petiole is greenish or yellowish and 5-10
cm long. The leaves are smooth, glabrous
and their margins entire. The flowers
are minute, monosexual and arranged in
paired rounded syconia in the leaf axils.
The syconium, with a diameter of 10-12
cm, become dark purple as they mature.
Introduction to Israel: Ficus
religiosa was introduced into Israel as
an ornamental at low altitudes in the
Mediterranean region. Due to its tolerance
for high temperatures it has also been
planted in gardens in arid areas of the
country.
Biological and Ecological Properties:
Ficus religiosa geminates and grows in
warm humid habitats in oases. The
trees become established and develop
in rock fissures and on cliffs near
waterfalls. Ficus religiosa is resistant to
high temperatures but sensitive to cold,
though less than Ficus benghalensis.
It reproduces by seeds and cuttings.
Pollination is by a tiny wasp
(Blastophaga quaraticeps) that invaded
Israel in the late 1960s. Observations
at the Ein Gedi Nature Reserve have
shown that the species is dispersed by
two native species, the bird, Tristram’s
grackle (Onychognathus tristramii) and
the fruit bat (Rousettus aegyptiacus).
These animals can disperse the seeds to
distances up to 15 km from the parent
tree. Ficus religiosa is a long-lived
species, and trees a thousand years and
older can be found in its natural range.
Distribution in Israel: Ficus religiosa
spreads to gardens and yards in the Tel
Aviv and Haifa areas. It has become
one of the most invasive tree species
in gardens and parks in the Jordan
and Yizre’el valleys. At this point Ficus
religiosa invades natural areas only in

the Dead Sea Valley, mainly in the Ein
Gedi Nature Reserve along the David
and Arugot rivers. The invasion in the
area began in the early 1980s, and Ficus
religiosa was the first of the three Ficus
species to invade. In the early 2000s
87 specimens, mainly seedlings and
saplings, were found along the Arugot
River. They originated from seeds from
a mere 15 trees planted in gardens in
the vicinity. Of the three invasive Ficus
species in Ein Gedi, Ficus religiosa is
the second most invasive after Ficus
benghalensis, even though its invasion
rate seems slower than that of the
latter. The large number of trees that
germinated in natural areas and the
distance from the seed source justify
classifying the species as “invasive” in
Israel.
Effect on Native Species: Ficus
religiosa, like other invasive species,
Geographic Origin: The natural range of Ficus
religiosa includes India, Pakistan, Bangladesh, Sri
Lanka, southern China and Tailand. In India F. religiosa
grows at altitudes up to 1500 m.
� References: 29, 30, 100, 118, 315.
can displace native species, particularly
those growing near streambeds. At this
time there is no information on the effect
Ficus religiosa has on the native fauna as
a result of local animal species feeding
on the syconia.
Control: Uprooting Ficus religiosa
seedlings is effective if the entire root
system is removed. It is not sufficient
to cut established and mature trees,
since the tree can renew itself. These
should be treated with specific chemical
methods, particularly hack and squirt
with triclopyr, which is considered very
effective against Ficus religiosa. No
biological control method has been
developed yet, nor does there seem
to be any development in the near
future. Effective management requires
119
uprooting all the trees in gardens in
the Dead Sea region. They can be
replaced by native tree species that have
equivalent scenic value and shade, but
do not threaten the natural vegetation of
the Ein Gedi Oasis.
Introduced Range: Ficus religiosa was
invasive for a time in southern Florida.
Seedlings were found in 1975 and in
1988, but they disappeared without
human intervention. Israel, mainly in the
Dead Sea Valley, is now the only place
in the world Ficus religiosa has invaded.
The tree is a popular ornamental
throughout the world. In light of its
invasion in Israel, there is a possibility
that the pollinating wasp will penetrate
other regions and consequently increase
the invasive sites of Ficus religiosa.
 Heterotheca subaxillaris (Lam.) Britton & Rusby
Camphorweed
120
Family: Asteraceae
Basic Description: Heterotheca
subaxillaris is a perennial branching
herbaceous plant with erect stems,
slightly woody at their bases. In its
natural range annual and occasionally
biennial specimens can be found. The
plant is generally 40-90 cm tall, but
some specimens may reach 1.5 m. It has
alternating lanceolate leaves, 2.5-10 cm
long and 0.8-3.5 cm wide. Both sides
of the leaf are rough, hairy and a bit
sticky. The leaf margins are crenulate
and the leaf surface is a bit wavy. The
leaves at the base of the plant are larger
than the upper leaves. They are slightly
elongated, with a short petiole and
encircle the stem. On the upper part of
the plant the stems branch into a panicle
with a number of yellow inflorescences,
18-22 mm wide. At the center of the
inflorescence are at least 20 disk flowers
surrounded by about 40 ray flowers. The
camphorweed has two types of achenes:
those that developed from the disk
flowers are about 3 mm long, triangular
and lack a pappus. The seeds from ray
flowers are longer and have a pappus.
In Israel Heterotheca subaxillaris blooms
from late spring to early autumn.
Introduction to Israel: Heterotheca
subaxillaris was introduced into Israel in
1975 in an attempt to stabilize moving
sands. A few seeds were planted on the
Galilee and Sharon coasts.
Biological and Ecological Properties:
The preferred habitat of Heterotheca
subaxillaris is coastal dunes and
disturbed areas such as roadsides.
The species is resistant to low
temperatures, particularly in its early
stages of development. Germination
can occur at temperatures as low as
30C. Experiments have shown that
leaf rosettes remain viable after being
exposed to temperatures of -150C for
four hours. The plant has particularly
long roots that penetrate deep into
the ground. Seedlings 1 cm tall, have
been found to develop roots 25 cm and
longer. Reproduction is via seeds. In the
United States one individual plant was
found capable of producing 5000-18,000
seeds, with 71% of the seeds viable.
Seed dispersal is by wind. A majority of
the seeds are dispersed to a distance
of about 15 m from the parent plant,
but some can reach distances as great
as 30 m. Most of the seeds from disk
flowers germinate immediately following
dispersal, whereas some two thirds of
the seeds from ray flowers germinate
about one year following dispersal. Some
of the seeds remain viable for over a
year, but there is no information on how
much longer they can remain viable.
A study carried out in Israel showed that
seeds buried below 4 cm in sandy soils
do not germinate. In agricultural areas
in the southeastern United States, where
Heterotheca subaxillaris is eruptive,
it can displace species of Conyza,
which indicates a very high degree of
competitiveness.
Distribution in Israel: In the 25 years
since it was introduced into Israel,
Heterotheca subaxillaris has invaded
the entire coastal sands along the
Mediterranean Sea shore from Ashkelon
up to the Galilee coast. The invasion
dynamics of this species indicates that its
minimum residence time is very short.
As early as 1990 Heterotheca subaxillaris
began spreading towards the southern
Coastal Plain. It invades roadsides
and disturbed habitats, but can also
invade and form very large and dense
populations on sand dunes that have not
been significantly disturbed. The species
has also invaded protected areas such as
the Bay Sands Reserve (south of Akko),
the Ma’agan Mikhael Coast Reserve on
the Carmel coastal plain, the Sharon 121
Coast National Park, the Nitsanim
Sands Reserve between Ashkelon and
Ashdod and many other protected areas.
Heterotheca subaxillaris is classified as
“invasive” in Israel.
Effect on Native Species: Heterotheca
subaxillaris competes with native species
for sandy micro-habitats, such as
depressions, and displaces other species
such as Artemisia monosperma. It
forms units with a high degree of cover
that stabilize sands and completely
transform the physical character of the
infested habitat. Future studies will
determine to what extent H. subaxillaris
affected the psammophilic animal
species that remained in the coastal
sands. The species is an exotic “habitat
transformer” species in Israel.
Control: Adult Heterotheca subaxillaris
can regenerate after being cut.
Therefore, mechanical control can only
be effective if the plant is uprooted when
young, in the rosette stage. At this stage
the root can be completely pulled out.
Chemical control is effective, especially
when the individuals are still at the
rosette stage. Spraying must be focused
in order not to harm other native
species. Herbicides such as paraquat,
122
 123

simazine and diuron have been found to Introduced Range: At present

be effective against the plant, yet their
application in natural environments may
be problematic. Since seeds buried at
depths exceeding 4 cm cannot germinate,
soil inversion may be an option, especially
in sandy soils, but this technique is
relevant only for restricted areas. There is
still no biological control method against
Heterotheca subaxillaris.
Heterotheca subaxillaris is considered
invasive only in Israel, and has not been
reported as an invasive in other parts
of the world. There is a possibility that
in the near future the camphorweed
will extend its invasion range northward
and spread to the Lebanese and Syrian
coasts.

Geographic Origin: The natural range of Heterotheca

subaxillaris in the eastern United States extends from
northern Mexico, via New Mexico and eastern Colorado,
north up to southeast Pennsylvania. Some scientists
believe that its natural distribution area includes the
entire southern United States, from coast to coast.

� References: 61, 144, 256, 302.

 Ipomoea aquatica Forssk.
Chinese Waterspinach, Water Spinach
124
Family: Convolvulaceae

Basic Description: Ipomoea aquatica
is a perennial species with trailing stems
with nodes 10-20 cm apart, from which
thin roots and erect leaves grow. The
trailing stems are very long (3-20 cm)
and the plant floats or roots in moist
banks. Its leaves are simple, glabrous,
arrowhead shaped , 3.5-17 cm long and
1-9 cm wide, with entire margins. Leaf
color is light green with a protruding
yellow central vein. Petioles are 3-14 cm
long. The flowers have a broadly funnel-
shaped corolla, with five longitudinal
folds, 4-7 cm in diameter. Flowers are
bright white, pinkish or lilac, usually
solitary, occasionally in inflorescences
with up to 5 flowers. In Israel Ipomoea
aquatica blooms in the summer. Its fruit
is an oblong or spherical capsule, 1 cm in
diameter, containing 4-6 seeds covered in
dense hairs.
Introduction to Israel: The first
record of Ipomoea aquatic in a natural
area in Israel was in 2009. Since then
it was reported in different regions in
the country. The plant was apparently
introduced into Israel by foreign
workers from Southeast Asia, who raise
Ipomoea aquatica for food. The Chinese
waterspinach is an edible plant, rich in
iron, and it is very popular in tropical
Asia.
Biological and Ecological Properties:
Ipomoea aquatica grows in freshwater
habitats such as shallow rivers and
marshes. It also spreads in drainage
ditches on roadsides and in irrigation
canals. Ipomoea aquatica grows in still
water, but unlike most other invasive
aquatic plants it can grow and develop in
flowing water as well. Plants take root in
moist soil on banks or in the shallow parts
of the water body. Ipomoea aquatica is
sensitive to cold and optimal development
requires temperatures higher than 24-
250C. It grows very rapidly, up to 10
cm a day. The plant reproduces mainly
asexually: it can develop roots at the
nodes along the trailing stem and recent
research suggests that stem fragments
with at least two nodes are necessary
for vegetative reproduction. When the
stem is cut its fragments are swept away
and take root in other sites. It can also
 Jordan valleys. In other areas invasions
will probably be seasonal and limited to
summer months. At this stage Ipomoea
aquatica has been classified as a “casual”
alien plant in Israel.
Effect on Native Species: Ipomoea
aquatica can interfere with water flow in
streams and canals. It forms dense mats
that create shade and displace native
plant species in the water and banks close
to the water line. Large concentrations of
Ipomoea aquatica slow down water flow
and facilitate mosquito reproduction.

Control: Manual uprooting is the

reproduce by seeds, and one individual
plant can produce 150-250 seeds, which
are dispersed mainly by flowing water.
The seeds do not germinate in water, and
most germination is seen on shores or
banks, near the water line.
Distribution in Israel: Ipomoea
aquatica was first recorded in a natural
area in Israel in September 2009 in the
upper section of the Harod Stream in
eastern lower Galilee. The plants took
root on the stream banks and in shallow
water within dense stands of Phragmites
australis. A short while later the plant was
found on the Na’aman River banks in the
Galilee coastal plain. In late 2011 it was
recorded in the Negev coastal plain near
the Gaza Strip. The plant’s requirement
for high ambient temperatures indicates
that it could form stabile populations in
moist habitats in the Dead Sea and the
most effective treatment, but requires
extensive time and care in collecting
all the rhizome fragments. Herbicide
treatment with 2,4-D was found to be
effective abroad, but this treatment in 125
water bodies or moist habitats could
harm native species and thus is not
recommended for natural water bodies in
Israel. No biological control method has
been developed yet. The most effective
and inexpensive treatment is prevention
by education and completely prohibiting
cultivation of the species in Israel.
Introduce Range: Ipomoea aquatica
is invasive in the Philippines where it is
considered the second most damaging
plant, particularly to agriculture. It is also
invasive in Texas and Florida; cultivation
in Florida has been legally prohibited
since 1973. Other invasive sites are found
in Puerto Rico, Hawaii and in most of the
Pacific islands.

Geographic Origin: The natural range of Ipomoea

aquatica is apparently in central China, particularly
in its southern part. In Southeast Asian countries the
species is cultivated for food, which makes it difficult to
determine its natural distribution area precisely.

� References: 16, 112, 128, 146, 189, 340.

 Lantana camara L.
Common Lantana, Largeleaf Lantana
126
Family: Verbenacea

Basic Description: Lantana camara is There is no information on when exactly it

an evergreen shrub up to 1.5-3 m tall, was imported into the country.
with many branches, some of which creep
Biological and Ecological Properties:
on the ground or climb on walls, trees
Lantana camara grows in sunny, moist
and other shrubs. It has grayish-brown
habitats. It possesses a well developed
stems and the young stems are covered
lateral and vertical root system and
in relatively sharp thorns. Leaves are 2-
grows in most types of soils, including the
10 cm long and 2-7 cm across, alternate,
poorest ones. Its growth rate is slow in
ovate and have a short petiole. They are
its early stages and rapid in later stages.
greenish-yellowish to dark green, with
Lantana camara reproduces by seeds
slightly toothed margins, their upper side
as well as asexually by root shoots. It
is rough and their underside is hairy.
produces flowers as early as the first year
Flower heads, whose diameter is 2-3 cm,
following germination. In moist habitats it
are borne on pedicels growing out of the
blooms throughout the year. In India it is
leaf axils. The yellow, pink and red flowers
pollinated by butterflies and moths, and in
are small, 4-8 mm across, and their color
Australia by bees. Its pollinators in Israel
changes with age. The fruit is a drupe
are unknown. The proportion of flowers
5-8 mm wide and contains a single seed.
that produce fruits and seeds is very high
At first the fruit is light green, and as it
and can reach 85%. Seeds are dispersed
ripens it changes to crimson and then to
to great distances by birds thus leading to
black. The fruits are usually in clusters
rapid formation of new concentrations. In
of up to 20 drupes joined together and
moist habitats the seeds can germinate
resembling raspberries. In Israel the
all year. Lantana camara has marked
species blooms throughout the year.
outstanding properties: its leaves release
Introduction to Israel: Lantana camara chemical compounds that are extremely
was introduced into Israel as a garden effective in preventing germination
ornamental and planted in many areas. and growth of other species. Studies
have even shown that some of these repeatedly in natural areas, particularly
compounds (lantanadene A & B) are in sites close to moist habitats, but also
effective in biological control of other in dry habitats. Lantana camara has been
invasive plants such as Eichhornia classified “invasive” in Israel. 127
crassipes. Common lantana regenerates
Effect on Native Species: A recent
after burning and cutting, and fires may
study determined that the main effect of
even increase the extent of infested
Lantana camara is due to its allelopathic
sites. It is considered one of the ten most
properties and competition with native
invasive plants in the world. In a list
species for soil resources. Lantana camara
published by the IUCN it was included in
concentrations displace native species due
the 100 most invasive organisms in the
to the large amount of shade created by
world.
the shrub thickets. The shrubs can grow
Distribution in Israel: Until recently
at a density of 5000 plants per hectare,
Lantana camara was not considered
as found in infested sites in Australia.
a species with invasive potential in
An Australian study showed that the
Israel, as it requires moist habitats
biodiversity of infested sites remained
for development and spreads mainly
stable if Lantana camara cover remained
in irrigated agricultural areas, such as
below 75%, but drops drastically
citrus grove on the Coastal Plain and
when the degree of cover exceeds this
date plantations in the Dead Sea region.
threshold. Moreover the fallen leaves of
Lately a number of Lantana camara
the plant are particularly rich in nitrogen
concentrations have been found in
that is released as the leaves decompose
natural, and sometimes even protected,
in the soil. There are no studies as
areas. Populations of the species have
yet on how this could encourage the
become established in the Coastal Iris
establishment of other invasive plants
Reserve in the southern coastal plain
that prefer high soil nitrogen levels,
and individual plants have penetrated
such as species of Acacia, Prosopis and
the Tel Yitshak Reserve in the Sharon
Parkinsonia.
region. A number of plants were found
in the Bet Yerah National Park (southern Control: Young specimens of Lantana
Lake Kinneret), and new foci have been camara can be uprooted, but other
located in the Ein Gedi and Perat River mechanical control methods are
reserves in the Judean Desert. The scope ineffective against adult plants that
of invasion is still limited, but during the have a well-developed root system,
past few years the species has been found as well as the ability to regenerate
128
following cutting. Effective herbicides
include glyphosate, triclopyr and 2,4-
D, but since this species invades moist
habitats, herbicides must be applied with
great care. There is no information on
focused chemical treatments that are
applied directly to the plant by drilling
or smearing. Implementing a treatment
regime that ensures a low plant density
level could be effective locally, but will
not prevent continued seed dispersal
to other sites. Many experiments have
been conducted as part of the process
to develop a biological control method
against the plant: 41 insect species that
feed off leaves or flowers of Lantana
camara have been released in at least 41
different countries in which the plant is
invasive. Only 26 of these insect species
became established. At this time, with
the exception of the Solomon Islands,
Hawaii and Guam (all situated in the
Pacific Ocean), biological methods have
not succeeded in controlling the species. A
number of scientists recently suggested
that insect species that attack the
root system could be more effective in
controlling the plant.
Introduced Range: Lantana camara
has invaded over 60 countries worlwide.
A number of countries, such as South
Africa, have forbidden planting and
have legislation requiring owners of
infested sites to eradicate the species.
The main invasive populations of
Lantana camara are found in southern
Europe, tropical Africa, South Africa, the
Mascarene Islands and the Seychelles in
the Indian Ocean, India and Southeast
129
Asia, Australia, New Zealand, the
southwestern United States and the
Galapagos Islands.

Geographic Origin: The natural range of Lantana

camara is tropical Central America, including the
Caribbean Islands.

� References: 42, 61, 105, 116, 132, 133, 149, 178, 192, 239, 294, 295, 310, 311, 327,
340, 356, 359
 Melia azedarach L.
Chinaberry tree
130
Family: Meliaceae

Basic Description: Melia azedarach is seeds were used to produce insect

a broad-leaved tree, 5-12 meters high.
Its central trunk is gray and furrowed,
whereas the younger branches are
smoother, reddish with purple spots and
patches. Its leaves are large, dark green,
20-50 cm long, bi- or tri-pinnate with 4-6
leaflet pairs. The leaflets are subdivided
into 3-6 secondary leaflets that are 5 cm
long and 2.5 cm wide. They are glabrous,
acuminate and have serrate margins.
The inflorescence in Melia azedarach is
a multi-flower panicle, with lilac flowers
possessing 5 sepals and 5 petals, 1-
1.3 cm long, and 10 stamens. Flower
diameter is 15-20 mm. In Israel Melia
azedarach blooms in April-May. The fruit
is a pale yellow drupe, 1-1.5 cm across,
with skin that wrinkles as it ripens. The
fruit contains 1-6 elongated seeds, 8-10
mm long and 7-6 mm wide.
Introduction to Israel: According to
reliable historic documentation, Melia
azedarach was already found in Israel
during the 16th century. The British
reported it as a common garden species
in the early 20th century. Its fruit and
repellant, oil and in folk medicine,
which could be the reason the tree was
introduced into the area several hundred
years ago.
Biological and Ecological Properties:
Melia azedarach grows in moist habitats
such as riverbanks, and in disturbed
habitats with high moisture levels, such
as the edges of cultivated fields, gardens
and roadsides. The species grows rapidly
and can reach a height of 8 meters
within 4-5 years. Melia azedarach grows
in most soil types and is resistant to
high temperatures and aridity. Its root
system is superficial and usually does
not extend deeper than 70 cm below
the surface. The diversity of the habitats
where the tree can grow testifies to its
unusual ecological plasticity. It flowers
a few years after germination and
produces a great number of seeds, which
have not, however, been quantified. The
tree reproduces from seeds as well as
vegetatively from root suckers. In Israel
its seeds are dispersed by fruit bats
(Rousettus aegyptiacus) and probably
also by other native animals. Seeds remain should be monitored. Melia azedarach is
viable for at least two years. classified as “invasive” in Israel.

Distribution in Israel: In addition to Effect on Native Species: The main

gardens, roadsides and disturbed areas,
Melia azedarach also invades a number
of nature reserves such as the Kare
Na’aman and Ein Nimfit reserves in the
Galilee coastal plain, the Hula and Ein
Bdolah reserves in the Hula valley and
the Ein Yehuda Reserve in the Jordan
Valley. In the Hula Nature Reserve the
tree also establishes itself within dense
and undisturbed reed stands. However
Melia azedarach generally forms only
small stands in these areas and therefore
the species cannot be regarded so far as
a major ecological threat. Nevertheless,
Melia azedarach has massively invaded
other areas with Mediterranean climates,
such as South Africa, and is able to invade
non-disturbed areas. This would suggest
that the dynamics of this species in Israel
131
effect Melia azedarach is displacement
of native plants. Its fruit is considered
poisonous but no cases of wildlife
poisoning have been recorded in Israel.
Control: Young specimens can be
uprooted. Cutting mature trees is not
effective because the tree regenerates
from the stump. Chemical treatment by
cut-stump or drill-fill with triclopyr is
effective. There are no biological control
methods against Melia azedarach.
Introduced Range: Melia Azedarach
is invasive in South Africa, including the
Cape Province, and has been banned
for planting and commerce. It is also
invasive in the southern United States,
particularly in Texas and Florida, as well
as in Hawaii.

Geographic Origin: Melia azedarach has an extensive

range and can be found from India to the tropical-humid
region in northern Australia, including Southeastern
Asia, China, Japan, the eastern Melanesian Islands and
the Polynesian Islands in the Pacific Ocean.
� References: 24, 107, 129, 149, 289, 293, 300, 340.
 Myriophyllum aquaticum (Vell.) Verdc.
Parrot’s Feather
132
Family: Haloragaceae
Basic Description: Myriophyllum
aquaticum is a perennial aquatic plant.
In most cases the species is dioecious,
but some individuals have been found
bearing both male and female flowers.
Most of the plant is submersed, but
compared to other species in this
genus, the tips of the upper leaves
in Myriophyllum aquaticum are erect
and can extend to a height of 15 cm
above the water surface. Its roots can
descend to depths of 2 m or more, in
order to reach the bottom. Myriophyllum
aquaticum has thin yellowish or reddish
stems, 2-5 m long and its leaves are light
green, 0.8-1.2 cm wide and arranged in
whorls of 4-6 leaves, 3-4 cm long. The
leaves are pinnate, with 10-14 pairs of
straight narrow leaflets, only 7 mm long.
The submersed leaves have fewer leaflets
than the emergent leaves. Myriophyllum
aquaticum has a small number of flowers
at the base of the leaf petiole, on the
emergent stems. The flowers are tiny,
only 1-1.5 mm long and lack petals.
Its fruits and seeds are unknown in the
areas the species invades.
Introduction to Israel: Myriophyllum
aquaticum was introduced into Israel as
an aquarium plant and is sold in plant
nurseries as a plant for ponds. The exact
time of introduction is not known.
Biological and Ecological Properties:
Myriophyllum aquaticum grows in
freshwater habitats such as ponds,
marshes, lakes and slow-flowing streams.
Only female specimens are found
throughout its invasion range where it
reproduces only asexually. Its stems
break easily, and a stem fragment only
5 mm long can develop roots and form
new plants. The species is spread only
anthropogenically, usually when aquaria
are cleaned out. Myriophyllum aquaticum
is resistant to cold: specimens have
survived for a year in temperatures of
+40C. When infested water bodies freeze
over the plants do not grow but remain
viable. The species can survive a lowered
groundwater level lasting up to 9 months,
provided the sediments remain saturated.
A recent Chinese study indicated the
species possesses allelopathic properties.
 plants. Thus Myriophyllum aquaticum
is considered an alien “transformer”
species in the regions it invades. The
species creates a favorable habitat for
mosquitoes. Its allelopathic effect on
native plant species has not yet been
studied.

Control: Removing Myriophyllum

Distribution in Israel: A small
population of Myriophyllum aquaticum
was found in Ein Nevoraya, a spring in
Biriya Forest (Upper eastern Galilee). The
plants covered the entire water surface
and ornamental fish were found in the
water, an indication that the origin of
the invasion was from the contents of an
aquarium emptied in the spring. Most of
the cases of aquatic plant invasions occur
in similar circumstances. In August 2011
the plant was found in the Alroee spring
in the Yzre'el valley in Lower Galilee.
In July 2012 it covered about 80% of
the spring's water surface. Most of the
biomass was mechanically removed
during summer 2012 (G. Gutman pers.
comm.). Myriophyllum aquaticum is
classified as a “casual” alien species in
Israel at this point.
Effect on Native Species: Myriophyllum
aquaticum forms dense invasive sites
that displace native plant species.
Their populations modify the physical
and chemical qualities of the water,
a trait found in many invasive aquatic
aquaticum from an infested water body
is a difficult task, therefore it is crucial
to prevent the formation of new foci.
Preventing the introduction of male
plants into Israel is essential in order
to avoid sexual reproduction. In South
Africa male plants were introduced in
the late 1960s and early 1970s and
brought about seed production which
rapidly resulted in an explosive increase
of new populations. Mechanical control 133
is effective only for a short time, since
there are always stem fragments that
remain in the water and form new plants.
The information regarding chemical
control is conflicting: according to some
sources 2,4-D and diquat are effective.
According to other sources chemical
treatments affect only the emergent plant
parts and do not eradicate the entire
organism, consequently the method is
not considered effective. However the
use of diquat was banned in Europe and
California in 2000 due to its harmful effect
on riparian ecosystems. At this point no
satisfactory method of biological control
has been developed, despite a number of
experiments conducted in South Africa.
Introduced Range: Myriophyllum
aquaticum is invasive mainly in Australia,
New Zealand, the western and southern
United States, South Africa and Great
Britain.

Geographic Origin: The natural range of Myriophyllum

aquaticum is tropical South America including Brazil,
Ecuador, Bolivia, Peru, Paraguay, northern Chile and
northern Argentina. According to some sources the
original range of Myriophyllum aquaticum was limited to
the Amazon Basin in Brazil.
� References: 32, 48, 85, 93, 127, 235, 239, 316, 340, 341
 Nicotiana glauca Graham
Tree Tobacco
134
Family: Solanaceae
Basic Description: Nicotiana glauca is
an erect evergreen shrub or small tree
with smooth stems and leaves, that
reaches a height of 4-5 m. Its leaves
are bluish-green and covered with a thin
layer of wax; they are alternate, elliptical
or oval, pointed and 5-25 cm long, with
the largest leaves up to 12 cm wide. Its
tubular yellow flowers (3-4 cm long) are
arranged in panicles at the tips of lateral
branches that hang down. The fruit
is an elliptical capsule, 7-10 mm long
and contains numerous seeds. In Israel
Nicotiana glauca blooms from June to
September.
Introduction to Israel: The first official
record of Nicotiana glauca in Israel is from
1898, although it is not clear how it was
introduced. It may have inadvertently
been brought in seed mixes or for use in
traditional medicine. Smoking tobacco
is prepared from a completely different
species that is herbaceous and not
invasive.
Biological and Ecological Properties:
Nicotiana glauca develops in disturbed
habitats such as roadsides, garbage
dumps and construction sites. In natural
areas the species mainly penetrates
along streams. In arid areas it is limited
to moist habitats such as streambeds.
The plant prefers habitats with deep
soils and high humidity. In its natural
range Nicotiana glauca is pollinated by
hummingbirds that feed off nectar, as
it also happens in California where the
species is invasive. In South Africa it is
pollinated by local sunbird species. In
Israel there is evidence that the Palestine
sunbird (Nectarinia osea) is active
around Nicotiana glauca flowers but its
pollination in Israel has not been studied
yet, thus it is unclear to what extent
local animal species are involved in the
process. An Australian study found that
new Nicotiana glauca populations may
establish themselves following unusual
flooding, and if there are no recurrent
floods the populations disappear after
a few years. The tree tobacco blooms
within a year of germination. Adult plants
can produce up to one million seeds
(in Australian invasive specimens). The
seeds are dispersed mainly by water,
and possibly also by wind. Seed viability
in the soil has not yet been studied. The
species possesses allelopathic properties:
a study from southeastern Australia
found that after the first seasonal rains
its leaves release chemicals that prevent
germination of annual species. Nicotiana
glauca renews itself rapidly following
cutting.
Distribution in Israel: Nicotiana glauca
is invasive in Israel’s Mediterranean
region, in the northern Judean Desert
and in wadi beds in the northern and
central Negev and in the Arava Valley.
The species invades roadsides and
garbage dumps, but also protected and
open areas throughout Israel such as the
Halilim Stream Reserve in the Jerusalem
mountains, Fault Escarpment Reserve
in the Dead Sea Valley, Wadi Besor
Reserve in the Negev coastal plain, the
Perat River and Wadi Og reserves in the
northern Judean desert and the banks
of Wadi Sekher in the Hatserim region in
the northern Negev. Nicotiana glauca is
classified as “invasive” in Israel.
Effect on Native Species: Nicotiana
glauca can form relatively dense stands,
135
which due to its large leaves, create
shade that displaces native vegetation.
Moreover the species’ allelopathic traits
affect native species germinating in its
vicinity.
Control: Nicotiana glauca seedlings can
be uprooted when they are still young. In
mature specimens cutting and spreading
glyphosate is effective but repeated
treatment may be necessary. There is
no biological control method against this
species.
Introduced Range: Nicotiana glauca is
a significant invader in southern Europe,
the Mediterranean islands, throughout
South Africa (where it has been declared
a weed), in Australia and the southern
United States – Florida, Texas, Arizona,
Nevada and California. The tree tobacco
is also invasive in Mexico, New Zealand,
Hawaii, the Canary Islands and the Cape
Verde Islands.

Geographic Origin: The natural range of Nicotiana

glauca is the tropical humid areas in western South
America, south of the Equator – Ecuador, Bolivia,
western Brazil, Peru, northwestern Argentina and
northern Chile.

� References: 61, 100, 110, 111, 116, 121, 130, 149, 246, 291, 302, 340.
 Oenothera drummondii Hooker
Beach Evening Primrose
136
Family: Onagraceae
Basic Description: Oenothera
drummondii is a perennial herb, slightly
woody at its base. Its branches are
prostrate and sprawl on the ground.
Oenothera drummondii is usually no taller
than 20-30 cm. Its branches and leaves
are covered by hairs that capture sand
particles flying in the wind. Leaves are
simple, slightly succulent, lanceolate, 4-
7 cm long and 1-1.5 cm wide. Oenothera
drummondii has bright yellow large
solitary flowers, 5-7 cm in diameter,
located on the tips of its sprawling
branches. They open at night and remain
open until the early morning hours.
Pollination is mainly by nocturnal moths.
The fruit is an elongated hairy capsule,
30-55 mm long and only 2-4 mm wide.
In Israel Oenothera drummondii blooms
from April to September.
Introduction to in Israel: The first
official record of Oenothera drummondii
in Israel is from 1912, but the species
was probably introduced in the late 19th
century, although it is not clear how. It
may have been inadvertently imported in
ships that came from the area of Houston,
Texas, as was the case in the 1920s in
southeast Australia. Another possibility
is that Oenothera drummondii was
introduced as a garden plant.
Biological and Ecological Properties:
Unlike the 15 other invasive species
of Oenothera in Europe, there is very
little information on the biological and
ecological properties of Oenothera
drummondii. It grows in sandy areas
along the Mediterranean shore, usually
at the head of dunes facing the sea, at
the edge of the vegetation cover. In these
conditions Oenothera drummondii forms
distinct non-continuous invasive foci.
The plant grows in regions with average
annual precipitation above 200 mm. It
reproduces only by seeds and has a deep
root system, whose characteristics, as
other traits of this species have never
been studied in depth.
Distribution in Israel: Oenothera
drummondii spread in coastal sands
along Israel’s coastline. It also invades
protected areas such as the Na’aman
Mouth, Ma’agan Mikhael Coast and
Poleg Stream reserves and the Sharon
Coast National Park. It has been classified
“invasive” in Israel.
Effect on Native Species: Oenothera
drummondii stabilizes dunes and can locally
displace native species. Its effect on the
coastal dune ecosystem in Israel is limited
as the plants do not form, at this time,
dense stands. Nevertheless, Oenothera
drummondii is an additional invasive
species characteristic of coastal sandy
areas and together with Carpobrotus edulis
and Heterotheca subaxilaris they are able
to modify the physical character of coastal
sandy habitats in Israel that are becoming
ever rarer in recent years.
Control: Uprooting young Oenothera
drummondii plants is possible, but may
prove difficult due to the deep roots.
Spraying herbicides is not advisable 137
as it may damage native species.
At this time there is no biological
control method against Oenothera
drummondii, nor does it seem probable
that any will be developed in the near
future.
Introduced Range: In the
Mediterranean Basin Oenothera
drummondii is invasive on the coasts
of Egypt, Morocco and Spain. It also
spreads along the southwestern and
southeastern coasts of Australia.

Geographic Origin: The natural range of Oenothera

drummondii extends from North Carolina to Texas, and
from there on to western and central Mexico.

� References: 61, 104, 162, 209, 246, 257, 302, 312.

 Oxalis pes-caprae L.
Bermuda Buttercup
138
Family: Oxalidaceae

Basic Description: Oxalis pes-caprae is a It was probably introduced into Israel

perennial herb with rhizomes that reaches from Europe.
a height of 15-30 cm. Its green leaves
Biological and Ecological Properties:
are composed of three heart-shaped
Oxalis pes-caprae grows in areas that
(chordate) leaflets with black markings
have an average annual precipitation
on their upper side. The petiole is 10-15
above 250 mm. It develops in relatively
cm long and extends directly from the
deep soils, in most soil types, including
rhizome buried in the ground. In darkness
sand dunes. Oxalis pes-caprae requires
the leaves fold backwards. The flowers
large amounts of light. In Israel it
are borne on an erect peduncle, 10-30 cm
reproduces exclusively asexually by
tall (taller than the leaf-bearing stems).
bulblets formed from rhizomes. A recent
The bright yellow flowers are trumpet-
study from the Balearic Islands showed
like with 5 petals fused at their base,
that one plant produces an average of
which open in the daytime and close at
three bulblets a year. Stems may develop
night. Oxalis pes-caprae develops from
from bulblets up to 60 cm below the soil
a white, hollow rhizome that is enveloped
surface. The bulblets are dispersed mainly
in thin layers of brown membranes. The
by people while cultivating a site, and also
rhizome is cone shaped and 2.5 cm long.
by birds and flowing water, mainly along
It lengthens creating small bulbs between
rivers and streams. Oxalis pes-caprae
the nodes. In Israel the species blooms in
invasive sites characteristically have large
January-April.
concentrations of plants, and densities
Introduction to Israel: The first record of 3000 plants per square meter are not
of Oxalis pes-caprae in Israel is from rare.
1906. The plant was imported into Europe
(Sicily) as an ornamental as early as 1796 Distribution in Israel: Oxalis pes-
and soon became an invasive plant in caprae populations have been found in
gardens, particularly in southern Europe. central and northern Israel, in the Yarkon
Sources National Park, the Acacia albida
Reserve near Ashdod, and in Einot Gibton
in the southern coastal plain, as well
as in the Coastal Iris, Bene Tsiyon and
Poleg Stream reserves located in the
Sharon. In northern Israel invasive sites
have been found at the Na’aman Mouth
and the Liman reserves located in the
Galilee coastal plain, as well as in the
Ammud Stream Reserve in upper eastern
Galilee. Oxalis pes-caprae is classified as
“invasive” in Israel.
Effect on Native Species: Oxalis pes-
caprae creates particularly dense carpets
that completely displace native geophytes
and annuals, and prevent development of
other herbaceous species.
Control: New small invasive
concentrations can be removed manually,
on condition that all the rhizome
and bulblet fragments are removed.
Uprooting is not effective; instead
it is necessary to dig up the ground 139
around the plant to be removed.
Chemical treatment with glyphosate is
very effective but does not completely
eradicate the plant. It is best to begin
treatment in early winter (November)
before the flowering season. There is no
method of biological control.
Introduced Range: Oxalis pes-caprae
is invasive in southern Europe and
around the Mediterranean Basin, mainly
in Portugal, Spain, southern France,
Italy, Greece, Cyprus, Syria, Lebanon,
Morocco, Algeria and Tunisia. The
species is also invasive in southeastern
Australia and in California.

Geographic Origin: The natural range of Oxalis pes-

caprae is restricted to the Western Cape Province in
South Africa.

� References: 18, 61, 125, 140, 183, 223, 239, 242, 302, 338, 340.
 Parkinsonia aculeata L.
Horse Bean, Jerusalem Thorn
140
Family: Caesalpiniaceae

Basic Description: Parkinsonia aculeata planted Parkinsonia aculeata in arid areas,

is an evergreen shrub, occasionally
a tree, 2-6 m tall, with crooked greenish
trunks that branch near the ground
and thin thorns up to 3 cm long. Its
leaves are pinnate, 20-40 cm long, with
small elongated leaflets (4-10 cm).
Flowers have 5 yellow petals, and their
diameter is about 2 cm. The upper petal
turns orange after the flower has been
fertilized. The flowers are arranged on
racemes with 8-12 flowers. Parkinsonia
aculeata blooms in Israel from April until
early autumn. The fruit is a light brown
thin pod, 3-6 cm long, occasionally up to
13 cm, and 7 mm wide, flattened between
the seeds. Seeds are olive or green, 9-10
mm long and 3-4 mm wide.
Introduction to Israel: Parkinsonia
aculeata was described by the British in
1922 as a sub-spontaneous tree. It may
have already been brought into Israel
during the Ottoman period. Parkinsonia
aculeata was used for afforestation
in sandy and arid habitats and has
become successfully acclimatized to local
conditions. The Jewish National Fund
e.g. north of Be’er Sheva, as part of its
savannization program in the late 1980s.
Biological and Ecological Properties:
Parkinsonia aculeata is very tolerant of
drought and cold. It grows in diverse
types of soil – sandy, saline, very acidic
(pH 3) and very basic (pH 11). The tree
grows very rapidly. Parkinsonia aculeata
reproduces by seeds only and a mature
individual can produce about 5000 seeds
annually. The seeds can float on water
and fall near the parent tree and are
dispersed by birds and water. Less than
1% of the seeds germinates in their first
year and most remain dormant in the
soil. There is no information on how long
they remain viable. Passage through
the avian digestive system improves
seed germination. Parkinsonia aculeata
regenerates following fire or cutting.
Distribution in Israel: Parkinsonia
aculeata invades disturbed habitats
such as roadsides and railroad tracks
and along other infrastructures as well.
Parkinsonia aculeata is also invasive in
protected natural areas, particularly in
moist habitats such as the Bet Tsayda
Nature Reserve in northern Lake
Kinneret, the Kare Na’aman Reserve in
the Galilee coastal plain, the Tavor River,
Yissakhar Stream, Jordan River Head-
Hamadiya and Arbel nature reserves in
the northern Jordan Valley where large
populations have become established.
According to Shmida (2005) Parkinsonia
aculeata became invasive only in the
early 1970s, about 50 years after it
was introduced into Israel. Parkinsonia
aculeata is classified as “invasive” in
Israel.
Effect on Native Species: Parkinsonia
aculeata forms dense groves that create
heavy shade that displaces all native
species.
Control: Uprooting large specimens is
effective, but requires monitoring as
new seedlings could develop at the site.
Chemical control is effective when applied 141
by drill-fill, cut stump or frilling methods
with 2,4-D combined with triclopyr. In
Australia an unsuccessful experiment
with biological control using the insect
Rhinacloa callicrates, which feeds on
P. aculeata leaves was conducted in
the 1990s, and at this time there is no
effective means of biological control.
Introduced Range: Parkinsonia aculeata
is one of the most problematic invasive
alien species in Australia and is included
in the WONS list. It is also invasive
in the western United States, Chile,
tropical Africa, South Africa, Hawaii and
Micronesia.

Geographic Origin: The natural range of Parkinsonia

aculeata is the southern United States, Mexico, the
Caribbean Islands, Central America and the northern
part of tropical South America.

� References: 14, 78, 239, 289, 302, 340.

 Paspalum distichum L.
Couch Paspalum, Knotgrass
142
Family: Poaceae (Graminea)
Basic Description: Paspalum distichum
is a perennial herb 15-60 cm tall. It
has creeping rhizomes and stolons that
branch on the soil surface. Its leaves are
narrow, rolled or flat, 3-13 cm long and 2-6
mm wide. The inflorescence is composed
of a pair of spikes (rarely one spike or
three) 2-7 cm long, located at the head of
the stem in the form of the letter V. The
spikelets are elliptical, 2.5-3.5 mm long.
Paspalum distichum blooms in Israel from
July to September.
Introduction to in Israel: The first
record in Israel is from 1939. Paspalum
distichum was introduced into Israel
unintentionally, presumably from the
northwestern Mediterranean where
Paspalum distichum spread after it was
introduced to France in 1802.
Biological and Ecological Properties:
Paspalum distichum grows in moist
habitats, e.g. riverbanks, lake shores
and marshes. It grows in a variety of
soils, with pH between 4.5 and 7.7.
Paspalum distichum is remarkably
tolerant of high heavy metal soil
concentrations. Reproduction is mainly
vegetative: fragments of rhizomes
develop roots within 36 hours, forming
new specimens. Growth rate is very rapid:
at temperatures of 300C a growth rate
of 3.3 cm daily was measured (1.8 cm
at 250C and 0.6 cm at 180C). Maximum
stable growth rate is at temperatures
between 300C and 400C. Plant fragments
are spread by floods and help the species
disperse to great distances in a short
time. It is unknown how long the seeds
remain viable. Paspalum distichum is
resistant to temporary flooding. Individual
plants can survive in water 50 cm deep,
as long as the upper section of the plant
is outside the water. Paspalum distichum
is also resistant to fire. This assortment
of properties indicates an unusual
ecological plasticity. Its only limitation is
its requirement for full light, as it cannot
grow in shade.
Ditribution in Israel: There are many
populations of Paspalum distichum in
Israel, including in protected sites: E.g.
In northern Israel – in the Hermon River,
Tel Dan, Hula and Ein Bdolah reserves
in the Hula Valley; the Yehudiya River
Reserve in the southern Golan Heights;
the Tavor River Reserve in the Jordan
Valley and the Ein Afek Nature Reserve
in the Galilee coastal plain. In the center
of the country the plant invades the Ahu
Binyamina and Poleg Stream reserves in
the northern Sharon, and the Einot Gibton
Reserve in the southern coastal plain.
Populations have also been found in the
Judean Desert seasonal streams, e.g.
Wadi Og . Paspalum distichum is classified
as “invasive” in Israel.
Effect on Native Species: Its combined
ecological properties allow Paspalum
distichum to form dense monospecific
swards that suppress tree seedlings as
well as herbaceous species and prevent
regeneration of native plants.
Control: Uprooting Paspalum distichum
is not effective unless it is possible
to ascertain all the roots have been
removed. Paspalum distichum does not
survive in the shade, so if field conditions
permit, it is possible to apply solarization 143
by covering the site with impermeable
polyethylene sheeting. However, this
method is difficult to implement near
water. Chemical control with glyphosate is
considered effective but cannot be applied
in wetlands. No biological control method
has been developed yet for Paspalum
distichum.
Introduced Range: Paspalum distichum
is invasive in all continents except
for Antarctica. Its main invasive sites
are found in southern Europe, Japan,
Southeast Asia (particularly Thailand),
Vietnam, Laos and the Philippines.
The plant is a constant problem in rice
fields. It is also invasive in India, Chile,
Argentina, New Zealand and southwestern
Australia.

Geographic Origin: The natural range of Paspalum

distichum is not clear, but it apparently ranges from the
southern United States to the Caribbean Islands and
tropical South America, from where it was introduced
into Western Europe. Some scientists consider it a native
species in northern Australia as well.
� References: 2, 3, 26, 61, 100, 152, 160, 162, 340.
 Pennisetum clandestinum Hochst. ex Chiov.
Kikuyu Grass
144
Family: Poaceae (Gramineae)
Basic Description: Pennisetum
clandestinum is a perennial plant, 30-40
cm tall and often taller, with branching
stolons, which attach to the soil surface
forming a dense mat. The base of the
leaves is yellowish or greenish, usually
with thin hairs. The leaves are long and
narrow, up to 30 cm long and 7 mm wide.
The young leaves are folded and open as
they mature. The inflorescence bears 2-
4 spikelets, 10-20 mm long, enclosed in
the upper leaf sheath. The spikelets are
sessile and bear two flowers. Seeds are
flat and black.
Introduction to Israel: Pennisetum
clandestinum was brought into Israel
as a lawn plant, apparently during the
British Mandate, but there is not sufficient
information relating to the specific date it
was imported.
Biological and Ecological Properties:
Pennisetum clandestinum invades moist
habitats such as riverbanks. It grows
on drained heavy and alluvial soils.
Pennisetum clandestinum can tolerate
soil acidity up to pH=4.5. Studies have
shown that peak photosynthetic activity
in Pennisetum clandestinum occurs at
temperatures between 250C and 400C.
The species is also frost resistant: the
leaves are affected at temperatures
of -20C, but clumps of the plant can
recover from exposure to temperatures
as low as -90C. It can tolerate brief
inundations, up to ten days, and is also
resistant to grazing and fire. Pennisetum
clandestinum is resistant to drought,
probably thanks to its root system:
although about 90% of its root volume
goes no deeper than 60 cm, some parts
penetrate to a depth of 5.5 m into the
soil. These properties are a sign of its
ecological plasticity. The plant requires full
light, and consequently does not invade
shady habitats. Reproduction is mostly
asexual via creeping rhizomes, and only
rarely by seeds. Pollination is by wind.
Distribution in Israel: A large
population, extending over 18 ha, has
become established in the Hula Nature
Reserve, located in the Hula Valley.
Small and medium-sized populations of
Pennisetum clandestinum are found in
numerous protected sites in Israel: Along
the Hermon River (Banias), in the Ein
Bedolah Reserve in the Hula valley, in the
Kare Ne’eman and Ein Nimfit reserves in
the Galilee coastal plain, in Timsah Pool
Reserve located in the Carmel coastal
plain. Pennisteum clandestinum invades
stream banks in the Judean Desert as well,
such as the Perat River Nature Reserve. It
is classified as “invasive” in Israel.
Effect on Native Species: Due to its
creeping rhizomes and mass on the
ground surface, Pennisetum clandestinum
smothers and displaces all other plant
species. Where it becomes established it
creates a dense lawn that is impenetrable
to native species. It may have allelopathic
properties but this has not yet been
unequivocally proven in field conditions.
Control: Small concentrations can be
uprooted, but it is important to ensure
that all root sections have been removed.
If conditions permit it is possible to apply
thermal disinfection by covering the
ground with impermeable plastic sheeting
for 4-12 weeks. Chemical treatment with
glypohsate is an option, but must be
accompanied by rehabilitation (planting
native species). Successful treatment of 145
Pennisetum clandestinum was carried
out in the Perat River Nature Reserve:
The foci distant from the river were
controlled with glyphosate (2%) while
the concentrations close to the river
were removed with imazapyr (1%). The
controlled patches were replanted with
Mentha longifolia, a native stream bank
species that creates dense coverage
and successfully prevents re-invasion
by Pennisetum clandestinum. In South
Africa biological control methods using
rust fungus was developed and in Hawaii
experiments were conducted with insects,
but up to now biological control methods
had only limited success.
Introduced Range: Pennisetum
clandestinum is an invasive species
in South Africa, Mexico, Brazil, Chile,
the Galapagos Islands, Australia, New
Zealand, Thailand, California and Hawaii.

Geographic Origin: The natural range of Pennisetum

clandestinum is in the humid tropical areas of East
Africa: Democratic Republic of Congo (formerly Zaire),
Tanzania, Burundi, Rwanda, Kenya, Ethiopia and
Uganda. In its natural range Pennisetum clandestinum
can be found at altitudes up to 2700 m.
� References: 100, 102, 126, 149, 162, 223, 237, 329, 340, 344.
 Phytolacca americana L.
American pokeweed, Pigeon berry
146
Family: Phytolaccaceae
Basic Description: Phytolacca americana
is a perennial erect, branching plant,
usually 1.2 m tall, but occasionally
reaching a height of 2.5 m. It has smooth
reddish-purple stems. Its leaves are
large, glabrous, lanceolate or elliptical,
10-13 cm long. The leaf margin is
entire. Phytolacca americana flowers are
arranged in drooping racemes, 15-25
cm long. Its flowers are white, 6 mm
in diameter. The fruit is a berry, with a
diameter of 9-10 mm when ripe. The fruit
ripens in late summer and becomes deep
crimson. Phytolacca americana blooms in
Israel from July through September.
Introduction to in Israel: The first
record of Phytolacca americana in Israel is
from 1898. It is not clear how the species
was introduced into Israel. It may have
been brought in as a medicinal plant or an
ornamental for gardens, as was the case
in Europe as early as the 17th century.
Biological and Ecological Properties:
There is little information on the
ecological properties of Phytolacca
americana. It has a thick taproot
(diameter up to 15 cm) that penetrates
deep into the ground and grows in
a variety of soils, heavy and sandy, with
a pH between 4.7 and 8. Phytolacca
americana grows rapidly and its seeds
are dispersed by birds, which unlike
other animals, are not sensitive to the
toxic substances in the fruit and other
parts of the plant. The plant is resistant
to cold and invades areas in Europe with
winter temperatures below -100C. In
Israel it can be found in low places, which
indicates that it is also tolerant of high
temperatures and can cope with a 7-8
months dry season. Phytolacca americana
invades disturbed areas such as grazing
areas and roadsides. Observations in
Israel have shown its ability to invade and
establish itself in dense natural vegetation
such as Rubus sanguineus thickets and
Phragmites australis stands, a trait that
indicates it has a high invasion capacity.
Distribution in Israel: Phytolacca
americana is invasive in the Hula Valley,
and mainly within the Hula Nature
Reserve, where it has massively invaded
a pasture of some 15 hectares, including
in a Rubus sanguineus thicket. Phytolacca
americana has been found in the Hula
Reserve regularly at least since the mid
1960s, but in recent years the extent of
its invasion has increased. Lately it was
observed in the Kare Na’aman Reserve
in the Galilee coastal plain. Phytolacca
americana is classified as an “invasive”
species in Israel.
Effect on Native Species: Due to its
strong allelopathic effects on local species
Phytolacca americana rapidly displaces
native plants in areas it invades. The
leaves possess chemical compounds that
prevent the germination of other plant
species. Thus, although the density of
Phytolacca americana stands in infested
areas is usually not very high the plant has
a strong negative impact on native plants .
Control: Uprooting Phytolacca americana
147
is not effective because the plant
regenerates. Removing the root system in
its entirety is very difficult. Concentrated
spraying of glyphosate is effective.
Treatment should be applied towards fall
and if necessary should be repeated. There
is no biological control method against
Phytolacca americana.
Introduced Range: Phytolacca americana
is invasive in most of the countries in the
Mediterranean Basin and in temperate
regions. There are reports of large invasive
concentrations in Portugal, France,
Hungary, New Zealand and South Korea.

Geographic Origin: The natural range of Phytolacca

americana is in the United States east of Kansas, and
in the southern United States from Texas to California,
Oregon and Washington state. The species also grows
in southeastern Canada.

� References: 28, 61, 100, 138, 143, 177, 206, 230.

 Pinus brutia Ten.
Calabrian pine, Turkish red pine
148
Family: Pinaceae
Basic Description: Pinus brutia is an
evergreen conifer, with a straight trunk,
20-25-m high, sometimes reaching 35
m. Its bark is grey-brown, with red-white
coloring in the fissures, even in young
specimens. The dark green, 10-15-cm long
needles are arranged in pairs and are 1-2
mm wide. The species is monoecious, with
male and female flowers (pine cones). The
male cones are small, about 2 cm long,
yellowish with red dots and the female
cones are large, ovoid and change their
color to purple and brown when they are
mature. At maturity cone length is about
11 cm and width is 4.5 cm. The cones have
a very short peduncle, are perpendicular to
the branch and are not slanted backwards
like in Pinus halepensis. In Israel Pinus
brutia blooms in April.
Introduction to Israel: Pinus brutia
was introduced into Israel in 1927 by
Jewish National Fund foresters for use in
afforestation. Extensive planting began in
the late 1980s.
Biological and Ecological Properties:
Pinus brutia is a hardy tree that grows
in most types of rocks and soils,
except for sandy areas. In its natural
range P. brutia grows in areas with an
annual precipitation between 350 and
1200 mm. It is more resistant to low
temperatures than the Aleppo pine and
can survive minimum temperatures
of -250C. Moreover P. brutia grows in
full light as well as in slightly shaded
conditions, which allows it to penetrate a
variety of vegetation units. For the first
10-12 years after planting Pinus brutia
grows faster than Pinus halepensis.
The calabrian pine is wind-pollinated
and can produce seeds 10 years after
germination. Its life expectancy is about
120 years, and occasionally can reach
150 years. Unlike Pinus halepensis it
is much less susceptible to the Israeli
pine bast scale (Matsucoccus josephi)
and for this reason it has been preferred
for afforestation over Pinus halepensis
during the past twenty years. Like most
pine species Pinus. brutia does not
renew itself after cutting or burning,
but its seeds have a high post-fire
germination rate.
Distribution in Israel: Pinus brutia
has been extensively planted only in
recent years; consequently there are still
only few infested sites. However, recent
but significant invasions of this pine
species have been observed particularly
within the native Tabor oak (Quercus
ithaburensis) wood parks, for example in
the Har Horshan Nature Reserve located
in the Ramot Menashe region. The species
is now classified as “invasive” in Israel.
Concern that the extent of its invasion
will grow in the forthcoming years is well-
substantiated, as the rate of new planting
has risen by some 50% in the late 1990s.
Effect on Native Species: Pinus brutia
completely modifies the structure and
composition of the Tabor oak wood park:
the pine establishes itself in the gaps
formed by the scattered cover of the oaks
where it creates a closed tree canopy
that prevents light from reaching lower
vegetation layers. Consequently the
herbaceous layer, which is particularly rich
in this type of wood park, is dramatically
reduced. Moreover, invasion increases the
risk of fire spread as the biomass of Pinus
brutia is extremely flammable. Dry pine
cones that remain on the branches ignite
rapidly bursting into flames; their burning 149
parts are carried to distances of several
meters and increase the speed of fire
spread in their vicinity.
Control: The only efficient treatment is
cutting. Since many planted plots border
the wood park and other protected
areas, control measures should be
implemented regularly to prevent Pinus
brutia concentrations from establishing
themselves and reaching maturity. In
view of the extent of planted areas in
Israel, and as a result of their proportion
in the country’s landscape, neutralizing
seed sources does not seem feasible in
the near future.
Introduced Range : Pinus brutia is not
known as an invasive species in other
parts of the world. In Australia it was
classified as a “naturalized” species in
2003. Unlike other pine species, Pinus
brutia was not planted in South Africa and
Chile.

Geographic Origin: Pinus brutia is a Mediterranean

species. Its natural range extends over northeast
Greece, Rhodes, Crete, west and southwest Turkey,
Cyprus and Lebanon.

� References: 7, 28, 138, 188, 208, 224, 225, 250, 258, 300, 302.
 Pistia stratiotes L.
Water Lettuce, Tropical Duckweed
150
Family: Araceae
Basic Description: Pistia stratiotes
is an herbaceous perennial floating
water plant, whose light green leaves
form a rosette. The leaves are fan-like
(flabelliform), up to 20 cm long and 8
cm wide. On the underside of its leaves
are six to eight prominent white veins.
The leaves are covered in short white
hairs that trap air and allow the plant to
float on the water. Its feathery roots are
yellow, thin and up to 80 cm long. Pistia
stratiotes is dioecious and has minute,
white unisexual flowers. Male and female
flowers are separated by a membrane.
The fruit is a caryopsis containing 14-
15 seeds each about 2 mm long. The
seed contains an air pocket. Its exact
blooming period in Israel is not known
but flowering plants have been observed
in early October.
Introduction to Israel: Pistia stratiotes
was introduced into Israel as an
ornamental pond plant and is still sold
as such in plant nurseries. There is no
information on when it was brought into
the country.
Biological and Ecological Properties:
Pistia stratiotes is invasive in tropical
and Mediterranean areas. Low
temperatures limit its invasion, but
studies have shown that seeds can
survive at least two months in water
temperatures of 40C and even for a few
weeks in ice at a temperature of -50C.
According to scientific literature Pistia
stratiotes can survive temperatures
between 150C and 350C following
germination, but population dynamics of
the species in the Hula Valley shows that
rosettes can cope with temperatures
as low as 100C. Reproduction is both
asexual and by seeds. When population
density is not high, reproduction is
vegetative. Stems about 60 cm long
in the water develop rosettes, which
then detach from the parent plant
and develop independently. When the
population density rises flowers develop.
Pollination is by wind and the seeds
mature after a month. For a while they
float on the water and then sink to the
bottom. The seeds germinate in light,
at temperatures over 200C, about 20-40
days after sinking. Pistia stratiotes seeds
can survive in the mud.
Distribution in Israel: Pistia stratiotes
has invaded a channel in the Hula Valley
and a small drainage channel near Yesud
HaMa’ala, in the same region, close to
the Hula Nature Reserve since at least
2004. The Hula Valley populations are
stable and they even survived the low
winter temperatures of 2007-2008.
There was a massive invasion of Pistia
stratiotes in the Alexander River, in
the northern Sharon, and the Lakhish
Stream, in the southern coastal plain, in
2005 and 2006, but these populations
did not survive. Winter flooding in the
Alexander River prevented permanent
invasion throughout the year, so that
damage was only seasonal. In late 2008
a new population was discovered in the
Tsalmon River about 50 meters from
Lake Kinneret. In summer 2009 isolated
individuals were discovered in the Timsah
Pool Reserve in the Carmel coastal plain.
In summer 2011 the plant was found
in the HaShofet Nature Reserve located
in the Ramot Menashe region. There is
concern that Pistia stratiotes could invade
Lake Kinneret, and were this to happen,
control will be long and complex. Pistia
stratiotes is still classified “naturalized”
in Israel, but as the number of invasive
sites grows, it will probably be classified
“invasive” in the coming years.
Effect on Native Species: When Pistia
stratiotes covers the water surface
completely it causes a drastic reduction
in dissolved oxygen and pH level. 151
Consequently, massive invasions of this
plant modify the abiotic character of the
aquatic ecosystem leading to the death of
most native aquatic organisms, making
Pistia stratiotes an ‘invasive transformer’
species. Dense layers of P. stratiotes also
block light from native aquatic plants
and displace them rapidly in infested
water bodies. In addition to ecological
consequences Pistia stratiotes also clogs
pumps and affects boating and fishing.
Invasive sites provide optimal conditions
for mosquito development.
Control: Manual or mechanical collection
of floating Pistia stratiotes plants is
effective in small concentrations, but
only if all the cleared plant are dried out
far from the water. Effective chemical
control has been achieved by spraying
the herbicide glyphosate (in formulations
suitable for aquatic habitats). 2,4-D has
been successfully tried, but treating
extensive concentrations is complex
and may not succeed in eradicating the
target population. Moreover, spraying
herbicides may lead to an accumulation
of organic matter in the water as a result
of the decay of dead plant biomass, which
could make rehabilitation of the infested
ecosystem even more difficult. A number
152

Pistia stratiotes in the Hula Valley in 2006

Pistia stratiotes in the Lakhish Stream in 2005 (southern coastal plain)

 153

of experiments have been conducted to Introduced Range: Pistia stratiotes

develop biological control methods with
various insects. At this point the most
successful treatment seems to be the
weevil Neohydronomus affinis, a native
of Brazil. Use of the weevil has had good
results in Australia, Texas and South
Africa. However recent experiments in
Florida have shown that the weevil feeds
of other plant species as well. Biological
control methods have suppressed target
populations but have not been able to
eradicate them completely.
invades mainly tropical and Mediterranean
regions such as South Africa, the
southern United States (particularly
Florida, Texas, Mississippi, Louisiana
and southern California), the Caribbean
islands, east and southwest Australia,
Hawaii and Turkey. It also invades many
areas in southeast Asia, such as southern
China, Cambodia, Indonesia, Thailand,
Malaysia and the Philippines, although
some experts feel that this is part of the
natural range of Pistia stratiotes.

Geographic Origin: The natural range of Pistia

stratiotes is not clear. Some scientists consider it a
pan-tropical species, whereas others believe it originates
in the humid tropical region in South America. This
assumption is based on the fact that a number of
insects and microorganisms that feed on Pistia stratiotes
originate in these areas.
� References: 5, 15, 49, 73, 79, 147, 149, 159, 162, 184, 239, 324, 340, 362.
 Populus alba L.
Silver-Leaf Poplar, White Poplar
154
Family: Salicaceae
Basic Description: Populus alba is a
winter deciduous tree, 20-30 m tall, with
a single, straight trunk. Young trees have
greenish-white smooth bark, which turns
grey-black at its base, with longitudinal
furrows, as the tree matures. Its leaves
are polymorphic: broad and ovate,
rounded or with triangular lobes. Leaf
length is 6-12 cm. the upper surface of
the leaf is glabrous and dark green, while
the lower surface is covered in dense
short hairs that give it a silvery white
hue. Populus alba is dioecious and its
inflorescence is a pendulous catkin. Male
catkins are hairy and their color is grey
and red. Female catkins are longer and
greenish. The fruit is a dehiscent capsule
with 2-4 valves. The seeds are small and
have wooly hairs that extend from the
base of the seed. In Israel Populus alba
buds and blooms in February and March.
Introduction to Israel: Populus alba
exists in Israel for over 2000 years and
may have been brought over by the
Romans. In those days it was customary
to plant the tree next to springs that
attracted many visitors, hence the name
Populus (meaning ‘people’ in Latin). In
the Middle East Populus alba was grown
near streams and springs because it
rapidly grows a straight trunk that can be
used for many purposes. Today Populus
alba is used as an ornamental in gardens
and is sold in tree nurseries.
Biological and Ecological Properties:
Populus alba grows exclusively on
river banks in the temperate and
Mediterranean regions and does not grow
in dry habitats. It can live for 300-400
years, grows in sandy or clay alluvial
soils and prefers basic soils. The tree has
a wide horizontal root system, grows very
rapidly, and forms large numbers of root
suckers. Pollination and dispersal are by
wind, but in Israel the species apparently
reproduces mainly vegetatively by root
suckers and can also regenerate from
stumps. Unlike other alien plants in
Israel, mature individuals of Populus.
alba are affected by a local species of
beetle Capnodis miliaris. The beetle larvae
develop in the roots and trunks of Populus
alba trees and form tunnels in the phloem
and the cambium. The beetle usually
attacks sick or dry trees and so cannot
eradicate entire populations.
Distribution in Israel: Populus alba
prefers moist habitats in cooler areas and
therefore is found in Israel only locally in
the Hermon region, particularly in the Tel
Dan and Hermon River (Banias) reserves,
where there is a large population of the
species. Populus alba seems to have
exhausted the spaces it is able to invade
in Israel but remains a management
problem in these two reserves. Populus
alba is classified “invasive” in Israel.
Effect on Native Species: Populus
alba forms dense units that create
extensive shade, which contributes to
the displacement of native species,
particularly those characteristic of river
banks. The trees that are destroyed by
the activity of Capnodis miliaris renew
themselves within a short time. Moreover,
the temporary gaps created by the
collapse of a mature tree damaged by the
beetle, accelerate the development of root 155
suckers and new saplings rapidly emerge
and replace the fallen trees.
Control: Repeated cutting may
neutralize mature trees. Chemical
control using drill-fill or cut and smear
are effective when performed with
glypohosate or triclopyr. Seedling leaves
can be sprayed with glyphosate (2%).
There is no available biological control
method.
Introduced Range: Populus alba has
been classified as an invasive species
in northeastern South Africa, the
northwestern and eastern United States
and in southwestern Australia (the Perth
region).

Geographic Origin: The natural range of Populus alba

extends from southwestern Europe (Spain, southern
France, Italy) via Eastern Europe to central Asia, up to
the Himalayas and central China.

� References: 100, 149 162, 260, 266, 285, 300, 340.

 Prosopis juliflora (Sw.) DC.
Mesquite, Devil’s Tree
156
Family: Fabaceae (incl. Mimosaceae)
Basic Description: Prosopis juliflora is
an 8-m tall tree with a broad flat canopy,
frequently with a number of trunks with
drooping branches that give it a shrub-like
appearance. The base of the petiole has
woody thorns up to about 5-cm long. The
leaves are bipinnate, 6-8 cm long, with
12-20 paired leaflets. The leaflets are
oblong, 5-12 mm long and 1.5-2.5 mm
wide, with rounded tips, glabrous upper
sides and slightly hairy undersides.
Prosopis juliflora has tiny, yellowish to
greenish flowers, borne on a tubular
raceme about 10 cm long. The fruit is a
brown pod with 10-20 oblong seeds, up to
7 mm long and 2-5 mm wide. In Israel the
species blooms from April to October.
Introduction to Israel: Prosopis
juliflora was introduced into Israel as an
ornamental and for afforestation. The
date of its introduction is unknown, but it
was probably after Israel’s independence.
The mesquite is still sold in nurseries
throughout the country.
Biological and Ecological Properties:
Prosopis juliflora invades along wadi beds
and roadsides in arid and semi-arid areas.
It is a hardy species that can tolerate
extreme soil salinity and aridity and grows
in both sandy and heavy soils on flat areas.
It can grow in water logged soils as well
as in dry soils. Its root system can reach a
depth of 15-20 m and also spreads laterally
close to the soil surface. Reproduction is
both by seeds and vegetatively by root
cuttings. Specimens can bear fruits after
3-4 years, occasionally later, depending
on habitat conditions. Recent studies have
shown that invasive seedlings in Mauritania
began flowering as early as two to three
months after germination. This indicates
that the biological development rate of
Prosopis juliflora outside its natural range
can change significantly and accelerate
the rate of invasion. Each tree produces
particularly large quantities of seeds.
Up to 2000 seeds/m2 were measured at
a soil depth of 9 cm in infested sites in
Kenya. The seeds are dispersed by running
water or by animals, such as goats, cattle
or camels that feed on the pods. Seed
passage in the animal digestive system
increases the germination rate. A study

carried out in Ethiopia showed that 1 kg
of goat droppings could have as many
as 760 seeds, of which 37% are viable.
Seeds can remain viable for 40 years.
Seedling viability is high due to the speedy
development of the root that penetrates
rapidly into the ground. The pods and
leaves contain allelopathic compounds.
Prosopis juliflora regenerates even after
severe wildfires and heavy grazing.
Distribution in Israel: Prosopis juliflora
is still limited to sites with a few individuals
in protected areas in Israel, i.e. the Ga’on
HaYarden Hamadiya Reserve in the Jordan
Valley and the Yotvata Reserve in the
southern Arava. In recent years new trees
have appeared in the northern Negev, i.e.
Wadi Besor Reserve and in the riverbed
of Wadi Ashan in the Negev coastal plain
where the tree has established dense
stands. Prosopis juliflora is now classified
as “invasive” in Israel. Its rapid invasion
rate requires intensified monitoring of the
development of new invasive sites in Israel.
Effect on Native Species: Prosopis
juliflora can create dense thickets that
displace native plants. Studies carried out
Geographic Origin: The natural range of Prosopis
juliflora extends from Mexico up to northern South
America (Ecuador and northern Peru).
� References: 8, 21, 25, 70, 90, 98, 162, 204, 222, 226, 239, 240, 241, 336, 340, 363.
in South Africa and in Kenya showed that
native plant diversity is significantly lower
in infested sites. The same applies for bird
species diversity in dense thickets formed
by the mesquite. Local plant species are
also affected by the allelopathic properties
of Prosopis juliflora, mainly by compounds
contained in the numerous pods that cover
the ground in the tree’s vicinity.
Control: Young specimens of Prosopis
juliflora can be manually uprooted. Cutting
alone is ineffective and may worsen the
situation as it stimulates the development
of root suckers. Local experiments in the
Negev suggest the cut-stump method with
application of triclopyr may be effective.
Spraying leaves with herbicides is not
effective. Attempts have been made to
develop biological control methods for 157
other Prosopis species, but none have
been developed for Prosopis juliflora.
Introduced Range: Prosopis juliflora
is the commonest invasive species in
its genus. It has invaded 120,000 ha of
arid land in Western Australia, where it
is included in the WONS list. In South
Africa it invaded 1.8 million ha of land,
particularly in the Northern Cape Province.
New data on the expansion rate of
invasive sites in South Africa’s arid region
indicate an annual invasion rate of 18%,
i.e. doubling the infested area within 4.5
years. The mesquite is a major plague
in Sudan, Kenya and Ethiopia where its
rapid encroachment on pasturelands has
led to tribal conflicts for the remaining
few gazing areas. In these regions the
mesquite is called the Devil’s tree. The
species has been declared invasive in
the United Arab Emirates, on the island
of Asunción in the Atlantic Ocean, on the
island of Réunion in the Indian Ocean and
in Hawaii.
 Ricinus communis L.
Castor Bean, Castor Oil Plant
158
Family: Euphorbiaceae

Basic Description: Ricinus communis is Jonah. Its oil (castor oil) is used in medicine
a shrub 1-4.5 m tall that branches from as a purgative. The first record of Ricinus
its base. It has very large palmate simple communis in modern Israel is from 1894.
leaves, with a diameter of 10-40 cm. The
Biological and Ecological Properties:
alternate leaves are a glossy dark green,
Ricinus communis is a pioneering plant
occasionally with red veins. The reddish
in moist habitats such as stream banks
trunk is smooth. Ricinus communis is
and marshes outside arid areas. It grows
dioecious and its flowers are monosexual
rapidly and can reach a height of 2 m within
and arranged in a 10-30 cm long panicle.
a year. It reproduces only by seeds. Seed
The minute male flowers are yellowish to
dispersal occurs by runoff, flowing water
greenish and their stamens have branched
and mud dispersed by animals. Ants are
filaments. The female flowers are located
also dispersal vectors as they collect the
above the male flowers and have three red
seeds to feed on a fat body at the head
pistils. The fruit is oblong, 10-20 mm long,
of the seed. Humans also disperse seeds
covered in soft spines and contains two
when transporting infested soils. The seeds
bean-like, brown seeds with black spots
remain viable for a number of years, but
that are 9-22 mm long and 3-4 mm wide.
no specific studies have been conducted on
In Israel Ricinus communis blooms from
the subject. The seeds can germinate and
March to November.
grow even when buried at a depth of 30
Introduction to Israel: There is no
cm. Seedlings can already produce seeds
information on when Ricinus communis
six months following germination. Ricinus
was introduced into Israel but there is
communis regenerates following cutting or
evidence it was used in ancient Egypt.
fire.
Ricinus communis probably grew in Israel
during the Second Temple Period, and Distribution in Israel: Ricinus communis
possibly even earlier. A plant bearing its is invasive mainly in Israel’s Mediterranean
Hebrew name is mentioned in the Book of region, particularly on stream banks and
other moist habitats. The southernmost
population of Ricinus communis is fund
in Wadi Besor Nature Reserve in the
Negev coastal plain, around the artificial
lake at the site. It is also common on
roadsides and less disturbed areas,
but there are also large populations in
nature reserves, i.e. the Hula Reserve in
the Hula Valley, the Ein Nimfit Reserve
in the Galilee coastal plain, the Ammud
Stream Reserve in the upper eastern
Galilee, the Yehudiya River Reserve in the
Golan heights, and along the Alexander
River in the northern Sharon. Medium-
sized populations are found in the Mikve
Udim reserve in the Sharon, the Kare
Na’aman Reserve in the Galilee coastal
plain, the Tavor River Reserve in the
northern Jordan valley, and in the Ein
Bedolah and Ein Ta’o reserves in the Hula
valley. Ricinus communis is classified as
“invasive” in Israel.
Effect on Native Species: Ricinus
communis forms dense units, particularly
in moist habitats. The extensive shade
it creates displaces native species
characteristic of these habitats. Its seeds
are extremely toxic to animals, but there
are only rare known cases of animal
poisoning. In the winter of 1969-70 a few
thousand ducks died in northern Texas
from Ricinus communis seed poisoning 159
and there has been evidence of leaf
aphids dying from ingesting the plant’s
phloem. It has also been found that
nematode populations decrease in areas
where Ricinus communis grows.
Control: Young specimens of Ricinus
communis can be easily uprooted, but
it is important to ensure most of the
roots have been removed, otherwise
the plant will regenerate. Concentrated
herbicide treatment using the cut stump
method with glyphosate (Roundup) or
triclopyr (Garlon), is effective if applied
correctly. Treatment is more effective if
implemented before fruits are formed.
There is no method of biological control.
Introduced Range: Ricinus communis
can be found on all continents except for
Antarctica. It is a large-scale invader in
the western United States at altitudes
below 300 m, as well as in southwestern
and southeastern Australia, Mexico, South
Africa (not including arid regions), Hawaii
and the Galapagos Islands.

Geographic Origin: The natural range of Ricinus

communis is not clear, but it is widely accepted that
it originates in tropical Africa. A number of scientists
believe that it also grows naturally in tropical Asia.

� References: 32, 61, 149, 170, 198, 239, 340.

 Robinia pseudoacacia L.
Black Locust, False Acacia
160
Family: Papilionaceae
Basic Description: Robinia pseudoacacia
is a winter-deciduous tree with a straight
trunk. Large specimens of the species
reach heights of 5-12 m in Israel. Young
trees have smooth trunks that become
dark gray with deep longitudinal furrow in
mature specimens. Its leaves are pinnate
with 7-20 elliptical leaflets (3 x 5 cm)
that end in a short point. Flowers are
white, 15-20 mm long and arranged in
drooping clusters 10-20 cm long. The fruit
is a straight flat brown pod, 5-10 cm long
and 1 cm wide, containing 3-10 black
seeds, 4-6 mm long. In Israel Robinia
pseudoacacia blooms in April.
Introduction to Israel: Robinia
pseudoacacia was described by the British
in 1922 as a sub-spontaneous plant in
Israel. It is planted as an ornamental
in public parks and along boulevards in
some cities in Israel.
Biological and Ecological Properties:
Robinia pseudoacacia grows in full
sunlight and develops in all types
of soil, with the exception of those
that are not well drained. Robinia
pseudoacacia is tolerant of cold and
drought. It reproduces by seeds as well
as vegetatively by root suckers. After the
first concentrations become established
the species continues invading, mainlly
by root suckers. Its growth rate is very
rapid, particularly in young trees (up
to 1 m annually). At the age of 6 years
the seedlings begin producing seeds,
but seed production peaks mainly 15-40
years after germination. Mature trees
produce numerous seeds every year or
two. Seed dispersal is by gravitation and
by wind. There is a large variability in the
time period seeds remain viable. Robinia
pseudoacacia absorbs atmospheric
nitrogen with the help of a symbiotic
bacteria that increases soil fertility. The
tree regenerates rapidly from stumps
after cutting or fire. Its life expectancy is
estimated at between 60 and 100 years.
Outside its natural range, e.g. in Europe,
trees 300 years old and older can be
found.
Distribution in Israel: Robinia
pseudoacacia does not form large
populations in Israel. A number of trees
have become established near a JNF tree
nursery that was once active in Kiryat
Anavim. In recent years several trees
have been reproducing at the edge of
the Judean Mountains National Park in
the Jerusalem Mountains region. Robinia
pseudoacacia has a high invasive potential
in areas with a Mediterranean climate and
therefore should be monitored in Israel.
At this stage Robinia pseudoacacia is
classified as “naturalized” in Israel.
Effect on Native Species: Robinia
pseudoacacia grows in exposed soil
pockets in concentrations of herbs and
shrubs, creating dense units that displace
native species that require light. Studies
have shown that Robinia pseudoacacia
concentrations prevent development
of pine species, due to the shade they
create. The trees raise the soil nitrogen
concentration and modify its chemical
balance. A recent study carried out in
Italy suggests that invasion by the black
locust, subsequent to the destruction of
native oak forests, was the major cause
of local extinction of native and rare
lichen species.
Control: Cutting Robinia pseudoacacia
161
is not effective, and can even have the
opposite effect, since the tree regenerates
from stumps and develops root suckers.
Chemical control using the cut stump or
drill-fill method with undiluted glyphosate
(Roundup) is very effective. The treated
infested area should be monitored for
10 years so new individuals that might
germinate from the soil seed bank can
be immediately eliminated. There is no
biological control method against Robinia
pseudoacacia.
Introduced Range: Robinia
pseudoacacia is invasive mainly in
California, Europe, Cyprus and North
Africa. It has recently begun to invade
South Africa (except for the northwestern
part of the country) and southwestern
Australia. Robinia pseudoacacia also
invades temperate regions of the world,
e.g. New Zealand, Argentina, Chile,
Canada, Korea and Japan.

Geographic Origin: The natural range of Robinia

pseudoacacia is the temperate region in the
southeastern United States in the area extending
between Indiana and Pennsylvania in the north, and
Louisiana and Georgia in the south.

� References: 32, 149, 162, 193, 199, 228,268, 277, 289, 300, 320, 340.
 Salvinia molesta D.S.Mitch.
Africa Payal, Giant Salvinia
162
Family: Salviniaceae
Basic Description: Salvinia molesta is
a perennial floating water fern, up to 30
cm long, with a short rhizome, which in
non-tropical areas may also be annual.
Salvinia molesta has two types of leaves:
submersed leaves, which are dissected
and serve as roots, and floating elliptical
leaves that are folded along the midvein,
1.5-6 cm wide. The size of these leaves
depends on the nutrient concentration
in the water. The leaves are covered by
a dense layer of hydrophobic hairs, 1 mm
long, that allow the plant to float. Salvinia
molesta populations create dense mats of
leaves between 2.5 cm and 1 meter thick.
Introduction to Israel: Salvinia molesta
was introduced into Israel as an aquarium
ornamental, probably sometime after
the 1970s. It was recorded in the wild in
Israel for the first time in July 2009.
Biological and Ecological Properties:
Salvinia molesta’s preferred habitat is
slow-flowing streams and water bodies.
It grows extremely rapidly: in subtropical
and Mediterranean regions an infested
area can double its extent in 2.2 days
in summer, and in 40 to 60 days in
winter. Population density can reach
1280 individuals per square meter. Its
growth rate is more rapid in early invasion
stages when the water body is still not
entirely covered. Optimal growth is at
300C, and the plant cannot survive more
than 2-3 hours at temperatures below
-30C or above 430C. The growth rate of
Salvinia molesta depends on the nutrient
concentration in the water, particularly
nitrogen in the form of the ammonia ion
[NH4]+. The species reproduces almost
exclusively asexually and only rarely by
spores. It disperses by water currents and
wind. The giant salvinia is known for its
effectiveness in removing surplus fertilizer
and heavy metals from water.
Distribution in Israel: A population
of Salvinia molesta was discovered in
the Yarkon River at the Yarkon Springs
National Park. The invasion in the Yarkon
River originated from aquarium water
emptied into the river. The plant was
found together with numerous Azolla
filiculoides plants and a few individuals
of Pistia stratiotes. At this point Salvinia
molesta is classified as a “casual” species
in Israel. The rise in the number of sites
with invasive aquatic plants in Israel
during the past five years and the fact that
this invasive fern also is sold in nurseries
in the country raises deep concern that
additional sites will be infested with
Salvinia molesta in the near future.
Effect on Native Species: Salvinia
molesta, like other invasive aquatic
species modifies the abiotic character of
the infested ecosystem, and is therefore
considered an “invasive transformer”
species in the areas it invades. The
species reduces pH and the concentration
of dissolved oxygen in the water and
displaces native aquatic species. Salvinia
molesta invasions have a negative effect
on water birds, fish and invertebrates that
require open water. Infested sites are also
excellent habitats for mosquitoes.
Control: As long as sites infested
by Salvinia molesta are small and the
invasion is in its initial stages mechanical
control methods are effective. The plants
can be manually collected with scoop nets.
For larger infested sites it is necessary to
use a mechanical shovel, but mechanical
methods can become complicated on
dense or unstable banks: due to the
relatively small size of the plant and its
fragility, rhizome fragments could scatter
in the water, so it is advisable to set up
floating barriers downstream from invasive
sites. In any case frequent monitoring for
long periods of time is necessary. Applying
herbicides to Salvinia molesta is not
always effective because the hydrophobic
hairs protect the upper sides of its leaves.
Moreover, chemical control of aquatic
plant species creates dead biomass that
accumulates in the infested water and
Geographic Origin: The natural range of Salvinia
molesta is southern Brazil.
� References: 38, 57, 95, 149, 215, 216, 223, 238, 239, 245, 319, 340.
releases large amounts of nutrients,
which in turn accelerates the renewal of
the invasion. For many years Salvinia
molesta was treated with diquat, which
is suitable for use in aquatic habitats, but
in 2002 the European Union banned the
use of this herbicide due to its negative
effect on river ecosystems. Diquat is
also banned in California. In the past
hexazinone and chlorosulfuron were
recommended to treat Salvinia molesta,
but in Europe the use of these herbicides
is prohibited in protected natural areas.
The aquatic herbicide fluridone was found
to be effective, but it too harms river
organisms and its use is forbidden in
Israel. The salvinia weevil, Cyrtobagous
salviniae, was found to be one of the
most effective biological control methods
against the plant. The treatment was 163
implemented successfully in Australia
in the early 1980s and since then in
a number of Southeast Asia countries, in
Africa and in the United States. However
a recent study carried out in Australia
suggests the weevil fails to persist over
the long term in some infested sites in
still water habitats.
Introduced Range: Salvinia molesta
invades many sites the world over:
In Africa it is invasive in the Zambezi
River, Congo River and Senegal River
in Mauritania and Senegal. It is also
invasive in Kenya and Zambia. In South
Africa it is classified as a weed. The giant
salvinia is invasive in Southeast Asia in
Sri Lanka, India, Malaysia and Indonesia.
It is also invasive in Australia (a WONS
species), New Zealand, Papua New
Guinea and New Caledonia. In the United
States Salvinia molesta invades the lower
Colorado River, between California and
Arizona.
 Schinus molle L.
Peruvian Pepper Tree
164
Family: Anacardiaceae
Basic Description: Schinus molle is an
evergreen tree, 5-12 m tall, with drooping
branches. It has pinnate leaves, up to 25
cm long, composed of 7-20 pairs of long
narrow leaflets (3-6 x 5-8 mm). At the
tip of the leaf is a single, smaller leaflet.
The leaf axis is not winged. Schinus molle
is usually dioecious and has monosexual
flowers. The flowers are arranged in
hanging panicles that bear several
dozen small white or yellowish flowers.
The female flowers develop pink to red
globular fruit with a diameter of 4-7 mm.
Each fruit contains a single seed. In Israel
Schinus molle blooms in July.
Introduction to Israel: Schinus molle
was brought to Israel by the British in
1919 as an ornamental plant. It was
planted in gardens and urban parks, and
occasionally along city streets.
Biological and Ecological Properties:
Schinus molle is extremely tolerant of
heat and aridity. It can germinate and
grow in dry habitats. A study conducted
in South Africa found that the habitat
preferred by Schinus molle seedlings is
the shade of local trees. Reproduction
is by seeds dispersed mainly by birds.
A recent Chilean study found that
mammals, including foxes, disperse the
seeds as well. The animals involved in
Schinus molle seed dispersal in Israel
have not yet been studied. The tree
possesses allelopathic traits: after its
leaves and fruit fall they release chemical
compounds that prevent germination of
other plants. This property makes the
tree very competitive. Schinus molle
regenerates rapidly following cutting or
burning. According to data from South
Africa, where the species is a problematic
invasive, the life span of Schinus molle
exceeds 200 years. Compared to other
invasive species the velocity of its spread
is medium to slow, so that neutralizing
invasive sites should be relatively easy.
Current Invasive Status in Israel:
Schinus molle grows in disturbed habitats
in Israel, such as roadsides, but has still
not formed large populations. It also
penetrates natural and protected areas
such as the Judean Mountains National
Park and the Halilim River Nature
Reserve in the Jerusalem Mountains,
where a number of trees grow.
Therefore, at this stage Schinus molle is
classified as “naturalized”. The species
was introduced into South Africa in the
mid 19th century but became invasive
there only in the 2000s, which indicates
that it has a long minimum residence
time, which extended over some 150
years. Therefore it would be advisable
to eradicate the few trees that have
become established in Israel, before they
become a source of seeds and cause a
widespread invasion.
Control Options: The effectiveness of
mechanical control of Schinus molle is
limited as the species regenerates following
cutting. Uprooting small seedlings may 165
be effective if most of the root system is
removed. Fire is not an effective control
method. Chemical control by drill-fill
methods could be effective, but there are
no specific details relating to recommended
herbicides. In order to improve the
effectiveness of controlling a given
population, chemical control should focus
on female plants. There is no biological
control method against Schinus molle.

Effect on Native Species: Schinus Introduced Range: Schinus molle is

molle affects native plant species, mainly invasive mainly in southeastern Australia
by its allelopathic properties. The soil and in South Africa. It is also invasive in
underneath Schinus molle canopies is California, Texas, Florida, Hawaii and the
completely empty of other plants. Galapagos Islands.

Geographic Origin: The natural range of Schinus molle

is in the semi-arid regions of northern Argentina and
Chile.

� References: 149, 167, 168, 214, 223, 289, 305, 306, 340
 Schinus terebinthifolius Raddi.
Brazilian Pepper Tree
166
Family: Anacardiaceae
Basic Description: Schinus
terebinthifolius is an evergreen shrub
or small tree, 1-7 m tall. It has pinnate
leaves, up to 8-15 cm long, composed of
3-4 pairs of elliptical leaflets (1-2.5 x 2.5-
7 mm). At the tip of the leaf is a single
leaflet. The leaf petiole is short, up to
3 cm long. The leaf axis is occasionally
winged. Schinus terebinthifolius is usually
dioecious and has monosexual flowers.
The inflorescence is 2-11 cm long and
has minute (1.2-2.5 mm) greenish-white
flowers. Male and female flowers look alike.
The female flowers develop pink or bright
red globular fruit with a diameter of 4-7
mm. Each fruit contains a single seed. In
Israel Schinus terebinthifolius blooms from
March to May.
Introduction to Israel: Schinus
terebinthifolius was brought to Israel
as an ornamental plant for gardens and
urban parks. The date of its introduction is
unknown.
Biological and Ecological Properties:
Schinus terebinthifolius grows well in
moist habitats near streams and marshes.
A recent study found that it reproduces
by seeds in moist habitats and by
root sprouts in dry habitats. A mature
individual of Schinus terebinthifolius
produces tens of thousands of seeds that
are dispersed mainly by native birds and
by water. There are still no studies on
which local bird species are involved in the
process in Israel. Studies have shown that
seed germination is high, particularly in
moist habitats, and that seedlings have a
high survival rate. Schinus terebinthifolius
seeds can become established and develop
in shady conditions. When the plant
overstory is removed, e.g. following a fire,
the seedlings grow rapidly (30-50 cm
annually). New individuals can produce
seeds three years after germination
but the seeds remain viable for only 5
months. Schinus terebinthifolius possesses
allelopathic properties: its leaves and fruit
release chemical compounds that suppress
germination of other plant species. The
trees regenerate very quickly following
cutting or burning. Schinus terebinthifolius
is included in the list of the 100 worst
invasive alien species in the world
compiled by the IUCN’s specialist group in
the field of biological invasions (ISSG).
Distribution in Israel: Schinus
terebinthifolius invades several areas in
Israel - the main populations are found in
the Ein Nimfit and Kare Ne’eman nature
reserves in the Galilee coastal plain, in
the Tel Saharon Reserve and in the Ein
Yehuda Reserve in the northern Jordan
Valley. Recently a population of Schinus
terebinthifolius was found in natural
brackish marshes in the Ne’ot HaKikar
area south of the Dead Sea where it
penetrates natural dense reed stands. In
Israel Schinus terebinthifolius has been
classified as “invasive”.
Effect on Native Species: In areas
Schinus terebinthifolius invades it forms
dense vegetation units with extensive
shade, thus displacing all the native
species. Its allelopathic properties are an
additional factor affecting native plant
species.
Control: Cutting is not an effective means
of controlling Schinus terebinthifolius as
it regenerates from stumps. Uprooting
small seedlings may be effective if more
than 75% of the root system is removed,
otherwise the plant will regenerate. Fire
is an effective control method for soil
seed banks, but mature trees regenerate
rapidly after fires. Chemical control
by drill-fill or cut-stump methods with 167
undiluted triclopyr is extremely effective.
In order to reduce the number of seeds
the treatment should be targeted at
female plants first. Herbicide spraying is
not very effective. Since the late 1980s
there have been attempts to develop
biological control methods, by using
the seed-eating wasp Megastigmus
transvaalensis, but they have not been
sufficiently successful. A study carried out
in Florida and published in 2011 showed
that biocontrol could be achieved with
seed rot fungi.
Introduced Range: Schinus
terebinthifolius is invasive mainly in
southwestern Australia, the eastern
part of the Kwazulu-Natal Province and
the Western Cape Province in South
Africa, central and particularly southern
California, Florida, Hawaii and the
Mascarene Islands in the Indian Ocean.
Schinus terebinthifolius is also invasive in
the western Mediterranean Basin.

Geographic Origin: The natural range of Schinus

terebinthifolius is in sub-tropical Brazil, Paraguay and
northern Argentina.

� References: 32, 78, 108, 149, 162, 296, 323, 340.

 Sesbania sesban (L.) Merr.
Common sesban, Egyptian pea
168
Family: Papilionaceae
Basic Description: Sesbania sesban
is a shrub or small tree, 1-7 m tall. It
has a central trunk with a diameter up
to 12 cm. The leaves are opposite, long
and narrow (4.5 x 18 cm), pinnate and
composed of 6-27 pairs of small oblong
leaflets (26 x 5 mm). The inflorescence
is up to 20 cm long and is composed of
2-20 yellow flowers. Sesbania sesban
blooms in Israel from April to the end of
August. The fruit is a straight or slightly
curved cylindrical brown pod, 20-30 cm
long and 2-5 mm wide that contains 40
seeds.
Introduction to Israel: It is not clear
how Sesbania sesban was introduced
into Israel, and there is only minimal
information on the topic. During the
1950s it was used as a windbreak in the
Jordan Valley and in the Ginosar Valley
located in the eastern upper Galilee
(Dan Pery, pers. comm.). According to
Dafni and Heller (1990) Sesbania sesban
“escaped” from gardens in the Jordan
Valley area. The date of invasion in Israel
is not known.
Biological and Ecological Properties:
There is little information on the
ecological characteristics of Sesbania
sesban. The plant grows in tropical
regions with one or two rainy seasons,
and in subtropical regions (including
Mediterranean climates). It grows rapidly
in all types of soil, including sandy
and saline soils. Its roots penetrate
deep into the ground. Sesbania sesban
grows in habitats that are flooded in the
rainy season and whose soil becomes
waterlogged. During floods it develops
floating roots. Aerenchyma tissue, a
spongy tissue with air cavities, protects
its trunk. Its seeds are viable for only
a short time, unless they are kept at
relatively low temperatures in a humid
environment. There is no information on
how Sesbania sesban disperses its seeds.
The plant has a short life span of about
5 years.
Distribution in Israel: The main area
infested by Sesbania sesban in Israel
is the Lake Kinneret shores where the
invasion began in 2007. A population

numbering hundreds of plants has
become established in the Kare Deshe
area (northwestern shore). Individuals
have reached heights of 2 m and
diameters of 6 cm. Solitary individuals
have also been found on the Ginosar
beach. Another population has become
established in the Bet Tsayda (Bteha)
Nature Reserve in northeastern Lake
Kinneret. Another large population is
known since 2002 from the banks of the
Sorek River near Kibbutz Tsor’a close
to Bet-Shemesh in the northern Judean
foothills. A few individuals were observed
on the Alexander River banks in the
Sharon. At this point Sesbania sesban
is classified as a “naturalized” species in
Israel.
Geographic Origin: The natural range of Sesbania
sesban has not yet been definitely determined, but it
probably originates in East Africa.
� References: 61, 80, 99, 141.
Effect on Native Species: Sesbania
sesban can form dense stands that
displace native species.
Control: Unlike most invasive woody
plants, Sesbania sesban is sensitive to
grazing and cutting, if the remains are
under 50 cm tall. Thus, cutting is an
effective mechanical control method.
Cutting is preferable to herbicide
spraying, whose effectiveness is
unknown. Several individuals in the Bet
Tsayda (Bteha) nature reserve were
controlled with the cut stump method
(cutting and applying herbicide) in late
summer of 2008, with apparent success.
There is no biological control method
against Sesbania sesban.
Introduced Range: Due to the
widespread use of Sesbania sesban as 169
a fodder plant throughout the world,
the species now grows all over tropical
Africa and in tropical and subtropical
Asia. The plant is also found in northern
and eastern Australia and in the Indian
and Pacific Ocean islands. The species is
not a significant nuisance in the natural
areas in those regions. Nevertheless,
Sesbania sesban has been classified
as a plant with invasive potential in
the United States and discovered as
an invasive species recently in central
Madagascar.
 Solanum elaeagnifolium Cav.
Silverleaf Nightshade
170
Family: Solanaceae
Basic Description: Solanum
elaeagnifolium is a perennial herb that is
usually 30-45 cm tall, but can occasionally
reach a height of 80 cm. Its stem and
leaves are covered in short dense hairs
that give the plant a silvery-white hue.
Its leaves are oblong with wavy ends,
6-10 cm long and 1-2 cm wide. Flower
diameter is 3.5 cm and each flower has
5 violet-lilac petals and 5 yellow stamens.
Solanum elaeagnifolium blooms in Israel
between May and August. Its fruit is
globular, smooth, 1.5 cm in diameter and
changes its color from green to yellow
and orange while ripening.
Introduction to Israel: Solanum
elaeagnifolium was first recorded in Israel
in 1957. There is no precise information
on how it was introduced into the country.
In other countries Solanum elaeagnifolium
was brought in mixed with other seeds for
agriculture or in fodder. The species was
probably introduced into Israel shortly
after the establishment of the state.
Biological and Ecological Properties:
Solanum elaeagnifolium grows in regions
with an average annual precipitation of
150-600 mm. It is resistant to drought
and to cold (up to -180C) and can thus
be found in semi-arid regions in all types
of soils, including sandy and saline soils.
The plant develops both a horizontal and
vertical root system. Some of its roots
penetrate to depths of up to 2 m, but
most of its root volume is fund close to
the surface, down to about 25 cm below
the surface. It reproduces both sexually,
by seeds, and asexually by rhizomes.
A root fragment one centimeter long
at a depth of 20 cm can regenerate
and produce a new individual. Ten-day
old seedlings are able to reproduce
vegetatively. There is disagreement as to
its main means of reproduction. According
to a number of studies the plant
propagates mainly via seeds and less via
its root system. Solanum elaeagnifolium
produces a large number of seeds: an
adult plant can produce between 1500
and 7200 seeds, which remain viable
for at least ten years. The seeds are
dispersed by wind, running water and
birds. Its fruits are toxic to domestic

animals. Allelopathic effects have been
reported.
Distribution in Israel: Solanum
elaeagnifolium invades primarily
cultivated fields but also adjacent natural
areas, among them protected sites e.g.
the banks of the Lakhish Stream in the
southern coastal plain, the Halilim Stream
reserve in the Jerusalem Mountains, and
the Tsippori National Park in the eastern
lower Galilee. The plant also invades
the arid region of the country as it was
recorded in the Wadi Besor Reserve in the
Negev coastal plain and in the Makhtesh
Be’eri Reserve in the southern coastal
plain. Solanum elaeagnifolium is classified
as “invasive” in Israel.
Effect on Native Species: Solanum
elaeagnifolium can create large and
relatively dense colonies primarily in
cultivated fields but also in adjacent
natural areas. Its extensive root system
usually spreads along the upper soil layers
and competes with native herbaceous
vegetation for water in the soil. Moreover,
it possesses allelopathic properties that
Geographic Origin: The natural range of Solanum
elaeagnifolium is the southwestern United States and
northeastern Mexico.
� References: 34, 35, 61, 94, 179, 239, 282, 292, 322.
provide it with a considerable advantage
over native herbaceous species. Solanum
elaeagnifolium can displace native
vegetation, particularly near water points
and at sites exposed to heavy grazing, as
livestock refrain from eating it.
Control: Solanum elaeagnifolium is
one of the most difficult invasive plants
to control and eradicate. It reproduces
rapidly from rhizomes, complicating
mechanical and chemical control
methods. Uprooting is not effective since
the roots break and the fragments that
remain in the soil allow the plant to
regenerate and continue multiplying. In
southern France an effective mechanical
control method was found to be uprooting
the whole plant with a large mound of
earth that includes the entire root system, 171
followed by burying the soil at a depth
of 3 m. Chemical control with picloram,
glyphosate or 2,4-D affects the foliage
but does not affect the root system and
is therefore ineffective in eradicating the
plants. Experiments with biological control
methods were conducted in South Africa,
Australia and the United States. A leaf-
eating beetle Leptinotarsa texana that
was released in South Africa in 1992 had a
significant effect on the foliage but did not
eradicate the plant. A nematode Orrina
phyllobia from Texas was successfully
used in Australia, but was found to
damage local Solanum species as well.
Introduced Range: Solanum
elaeagnifolium is invasive in all South
African provinces, Southern Australia,
India, Taiwan, the Mediterranean Basin
countries, particularly Morocco and
Greece, the central and eastern United
States, Hawaii, Argentina, Brazil, Chile,
Paraguay and Uruguay.
 Verbesina encelioides (Cav.) A.Gray
Crown Beard, American Dogweed
172
Family: Asteraceae (Compositae)
Basic Description: Verbesina encelioides
is an erect annual, 60-120 cm tall. Its
stem and the underside of its leaves are
covered in short, dense white hairs. The
leaves are 4-10 cm long and 2.5-6 cm
wide. The leaves at the base of the stem
are opposite and triangular, while on the
upper part of the stem they are alternate
and hastate. Leaf margins are toothed.
The inflorescences are bright yellow,
chrysanthemum-like and 2.5-5 cm in
diameter. In Israel the species blooms
from April until the summer. The seeds are
winged and have a pappus with two awns.
Introduction to Israel: Verbesina
encelioides was first recorded in Israel
along the Coastal Plain in 1970. In 2001 it
was discovered in the northwestern Negev.
It most probably was introduced into Israel
in seed mix shipments for agriculture.
Biological and Ecological Properties:
Verbesina encelioides is extremely
drought resistant and can grow in regions
with 150-200 mm average annual
precipitation. It grows in sandy, but not
saline soils, or in direct sunlight. Verbesina
encelioides reproduces only by seeds
that it produces in very large numbers:
one inflorescence can produce 300-350
seeds annually. Each plant can develop 2-6
inflorescences, so that its seed production
potential is between 600 and 2100 seeds.
The seeds are dispersed mainly by wind,
but due to the relatively large seed mass,
dispersal distance is not large. Recent
observations in the Negev desert showed
the seeds are also dispersed by native
ant species. The short awns at the tip
of the seeds stick to animal coats and
thus are dispersed to greater distances.
The seeds remain viable for long, but
there is no information on exactly how
long. Experiments conducted in Israel
showed that the germination rate
can reach 92%. Seeds on the ground
surface have a greater germination rate.
Germination has been observed from
seeds up to 7.5 cm deep. They germinate
at 5-350C and maximum germination
occurs at temperatures between 10 and
200C. According to this same study the
determining factor in the distribution of
Verbesina encelioides in Israel is soil type.

Most of the invasive sites in Israel develop
on sandy or loessial soils. The plant can
be very toxic to animals and therefore is
not eaten by them. Verbesina encelioides
has allelopathic properties that prevent
other plants from germinating.
Distribution in Israel: Small
populations of Verbesina encelioides can
be found in the Sharon near cultivated
fields, and from there the species invades
natural protected areas, such as the
Kadima Reserve. A larger population,
composed of a number of sites is found
in the Wadi Besor Reserve in the Negev
coastal plain, and in the Agur sands
area in the northwestern Negev. In this
area Verbesina encelioides exhibits two
patterns of invasion: the first on the
edge of agricultural areas, where dense
populations have become established;
the second via scattered individuals that
penetrate natural undisturbed areas in
the direction of the Agur Sands Reserve.
The ability of the crown beard to invade
Geographic Origin: The natural range of Verbesina
encelioides is in Mexico and the southern United States.
� References: 61, 89, 96, 103, 239, 262, 288, 298, 302.
undisturbed sites, far from agricultural
areas, indicates a particularly high
invasive ability and raises concern that
the species will continue to spread
into the Agur Sands Reserve. Recently
a large population of the plant was
found in a loessial plain southeast of the
Holot Mash’abim reserve in the eastern
part of the Agur sands area. Verbesina
encelioides is classified as “invasive” in
Israel.
Effect on Native Species: The shade
created by dense units of Verbesina
encelioides and its allelopathic properties
displace other plant species. Verbesina
encelioides is toxic to mammals.
Control: Removing flower heads before
seed production and preventing seed
accumulation on the ground is the 173
primary control method in small invasive
sites of Verbesina encelioides. Individual
plants can also be uprooted manually,
but the effectiveness of this treatment
is contingent on removing all the roots.
Chemical control by herbicides, a mixture
of glyphosate and triclopyr in dense
units, or glyphosate alone in sites where
Verbesina encelioides grows with native
species, is also a possibility. There is no
biological control method against this
species.
Introduced Range: Verbesina
encelioides is an invasive species mainly
in eastern and southwestern Australia,
northern India, Morocco and Midway
Island (Hawaii). Other sites exist in
the southwestern Arabian Peninsula,
Argentina, Namibia, Botswana and South
Africa.
 Washingtonia robusta H. A. Wendl
Mexican Fan Palm
174
Washingtonia filifera (Lindl. ex Andre) H.A.
California Fan Palm
Family: Palmae
Basic Description: The genus
Washingtonia is represented in Israel by
two species W. filifera and W. robusta.
Washingtonia species are dioecious, i.e
male and female flowers are on separate
trees. Both species are very similar and
difficult to tell apart in their early stages
of development. As in most palm species
both Washingtonia species have straight
trunks with no branches. The canopy
is composed of large leaves, 1.5-2.5
m wide, arranged like a fan. In Israel the
species blooms between March and May.
Washingtonia filifera is a medium-tall
tree, with a wide trunk, whose diameter
can reach 100 cm in mature specimens.
Its leaves are erect and droop a bit at the
edges. Washingtonia robusta is a tall tree
with a narrower trunk, whose diameter in
mature trees does not exceed 80 cm. Its
leaves droop. The lower side of the base
of the frond is covered with felt-like light
brown hairs. There are hybrids between
the two species, sometimes referred to
as Washingtonia x filabusta, that exhibit
intermediate traits.
Introduction to in Israel: The first
specimens of Washingtonia robusta
were brought into Israel in 1905
as ornamental trees. Washingtonia
filifera was introduced at a later date.
Sometimes both species were planted
together.
Biological and Ecological Properties:
Washingtonia trees are resistant to heat
and aridity. Washingtonia robusta is
more sensitive to cold than W. filifera.
In tropical and subtropical regions
that they invade the trees establish
themselves in a variety of habitats,
moist as well as extremely arid. Another
characteristic indicative of non-specific
ecological requirements is the species’
ability to germinate in shallow, poor
soils: Washingtonia trees can even grow
in the cracks between pavement stones.
Reproduction is only via seeds that are
 sites have been observed and we can
consider the species is still in its lag
phase. For these reasons Washingtonia
species in Israel should be classified as
“naturalized”. However, the increase in
the number of reports of Washingtonia
invasions from other parts of the world
in recent years, e.g. Southwest Australia,
California, suggests a follow-up of these
species in Israel is necessary.

Effect on Native Species: Washingtonia

usually dispersed by birds that spread the
seeds to distances of dozens of meters
and more.
Distribution in Israel: Washingtonia
robusta seems to be the more invasive
species of the two. It invades mainly
in gardens and roadsides. Lately trees,
mostly young (up to 4 m tall) have been
found in nature reserves and natural
areas. In the Mouth of the Na’aman
River Nature Reserve in the Galilee
coastal plain about a dozen W. robusta
trees have become established. In
the area of the mouth of the Tsalmon
River on the shore of Lake Kineret
(Sea of Galilee) young specimens were
found. A few trees, probably W. filifera,
established themselves in the Ein Gedi
Nature Reserve in the Dead Sea Valley.
Individuals were also reported in the
coastal sand dunes south of Tel-Aviv.
The number of invasive trees in natural
areas in Israel is still limited. No dense
species can outcompete other plants
thanks to their dense root masses,
and by creating shade. A recent study
showed how native arthropod abundance
steeply decreased in habitats invaded
by Washingtonia filifera. Moreover,
the accumulation of dry leaves along
the trunk increases the danger of fires 175
breaking out.
Control: The most effective treatment
is uprooting young specimens. Cutting
mature trees at their base is also
effective. Lately there has been success
in controlling mature specimens, in the
southwestern USA, using drill-fill with
glyphosate (50%); imazapyr is preferred
in wetland and riparian zones. There is no
biological control method.
Introduced Range: Washingtonia
robusta is invasive in Florida, southern
California and in Hawaii. W. filifera has
recently become invasive in the Perth
area in southwest Western Australia and
also in isolated desert springs in the
Death Valley National Park (California
and Nevada,) where it displaced native
shrubland vegetation.

Geographic Origin: The natural range of Washingtonia

species is the arid and semi-arid areas in the Sonora
desert in the southwestern United States and in northwest
Mexico. The trees grow at sea level along canyons and
near water sources. W. filifera is found in southeastern
California and in western Arizona, whereas W. robusta
grows in more southern and hot areas in northwestern
Mexico and in the Baja California peninsula.
� References: 158, 162, 175, 234, 302
 Xanthium strumarium L.
Common cocklebur, Rough cocklebur
176
Family: Asteraceae (Compositae)
Basic Description: Xanthium
strumarium is a perennial herb 30-120 cm
tall. Its leaves are triangular, irregularly
lobed and have a long petiole (2-8 cm);
they are alternate, 4-12 cm long and
covered with short hairs on both sides.
The species is a monoecious plant with
small green flowers. Male inflorescences
are located above female ones, and are
5-8 mm in diameter. In Israel the species
blooms in the summer months. The fruit
is rounded and oblong, up to 2 cm long.
Its surface is covered in hooked thorns
and on its apex it has two straight or
hooked bristles; as it ripens the fruit
turns from green to yellow, and finally
to brown. Fruits are borne in clusters of
up to a dozen and each one contains two
brown or black seeds.
Introduction to Israel: Xanthium
strumarium was first found in Israel in
1921, but it is not clear how it reached
the area. Possibly its seeds were
imported inadvertently together with
seeds intended for agriculture. Another
possibility is that Xanthium strumarium
fruits were brought in with sheep wool
imported to Israel from Australia or the
United States.
Biological and Ecological Properties:
Xanthium strumarium grows in moist
habitats, particularly riverbanks, seasonal
pools and near other water bodies. The
species has a well-developed root system
and grows rapidly, properties that increase
its competitiveness. Xanthium strumarium
can produce seeds at a very early stage in
its life, when it is only a few centimeters
tall. Dispersal is by animals that carry the
fruits on their coats. Moreover the fruits
have air pockets that help them float on
the water, allowing the plant to invade
rapidly along streams. The lower seed
in the fruit germinates in the spring or
summer after it has ripened, whereas the
upper seeds remain dormant for two or
three years, and occasionally even longer.
Distribution in Israel: Xanthium
strumarium grows on stream banks and
seasonal pools in Israel’s Mediterranean
region and in the Judean Desert wadis.
It is particularly common along polluted
streams such as the Lakhish Stream and
the Alexander River. The species has also
invaded Wadi Og and the Perat River
reserves. Xanthium strumarium is invasive
in the Hula Reserve in the Hula Valley
and around Lake Kinneret and in the
Btekha and Tavor River reserves. Other
populations exist on the Galilee coastal
Plain, such as in the Ein Afek, Ein Nimfit
and Kare Na’aman reserves. The plant is
found also in the Brekhat Ya’ar and Poleg
Swamp reserves. To the south, large
populations have established themselves
in seasonal pools such as the Zmorot Pool
and Pura nature reserves in the Negev
coastal plains. Xanthium strumarium is
considered an “invasive” species in Israel.
Effect on Native Species: Xanthium
strumarium can create very dense
stands, up to 30 plants per square meter,
that displace native species growing on
banks of rivers or other water bodies. It
is possible the species has allelopathic
properties, but this has not yet been
studied.
Geographic Origin: The natural range of Xanthium
strumarium is the western United States.
� References: 61, 239, 340.
Control: Small invasive sites of
Xanthium strumarium can be uprooted
manually. Chemical control using D-2,4
is effective, but only if sprayed before 177
the plant blooms. Burning may be
effective if sufficiently high temperatures
are achieved. Experiments with biological
control began in the 1930s in Australia,
but success has been achieved only since
the 1980s, when Epiblema strenuana,
a type of gall moth, was introduced to
Australia in 1982. In the late 1990s
Puccinia xanthii, a rust fungus was
used to suppress Xanthium strumarium
invasions in Queensland, Australia, but
in recent years the fungus has begun
attacking sunflower crops.
Introduced Range: Xanthium
strumarium is considered a particularly
problematic invasive species in 28
countries, including Australia, South
Africa, India, China, Japan, Korea, the
southeastern United States, Hawaii and
countries in the Mediterranean Basin.
178
 Glossary

Allelopathy The phenomenon in which a plant excretes chemical substances via

its leaves after it has shed them, or via other parts of the plant (e.g.
roots or seed coats), which prevent germination of seeds belonging
to other plant species at the site. Allelopathy is the main means of
direct competition between plant species and is relatively common
in invasive plant species. Allelopathic properties in invasive plants
significantly increases the damage caused to native plant species.

Blade The broad portion of the leaf.

Bulblet A small bulb produced for reproduction that develops at the base of
the mother bulb.

Capsule A dry fruit with multiple seeds that opens when it matures and
propagates its seeds.

Dioecious A plant with monosexual flowers (either male or female) where 179
the female flowers and male flowers are on separate plants. This
property can be advantageous when dealing with invasive plants, as
it is sufficient to neutralize individuals with female flowers to prevent
seed production in the treated population

Drupe A fleshy or succulent fruit with a stone enclosing a seed.

Evergreen A plant that has green leaves throughout the year. The leaves are
replaced and renewed individually and gradually, unlike winter or
summer deciduous plants.

Fern Perennial herbaceous species, flowerless, reproducing via spores,

that belong to the Pteridophyta,

Head (capitulum) An inflorescence with dense multiple flowers on short stalks or on

a receptacle forming a spherical or ovate structure. The invasive
Australian acacias in Israel have spherical heads.

Inflorescence A cluster of flowers on a stalk, a number of branches or an entire

plant that appear as one unit.

Monoecious A plant with androgynous flowers or monosexual flowers (male and

female flowers) on the same plant.

Morphology The study of form – a term relating to form, structure and appearance
of organisms.

Panicle An inflorescence with a principal axis that branches out into flower-
bearing branches.

Perianth The outer leaves of the flower that envelop the reproductive organs
(anthers and stamen).

Petiole The narrow part of the leaf that is connected to the plant branch on
one side and to the blade on the other.
 Phyllode A broad petiole that functions as a leaf. Some of the invasive
Australian acacias in Israel, such as Acacia saligna, bear small,
pinnate leaves for about two months following germination, which
they subsequently shed. These are replaced by simple phyllodes that
function as leaves.

Polymorphism A property that indicates the existence of diverse forms, structures

or appearance in one organism.

Raceme An elongated inflorescence with a principal axis bearing flowers on

pedicels.

Receptacle The broad end of the flower axis that bears the floral organs.

Seed viability The period of time in which seeds can germinate and form a new
individual. In many invasive plants plant viability is preserved for
a long period after the seeds have matured and been dispersed.
Therefore, to eradicate invasive plant populations it is not sufficient
to neutralize seed-bearing plants. Regular monitoring should
continue for the entire period the seeds remain viable. Often the
length of this period is not known.
180
Sepals The outer leaves of the perianth, usually green and not colored
(unlike the petals)

Sheath The broad base of a leaf that envelops the plant stem. Characteristic
of many species belonging to the Gramineae or Liliaceae families.

Spike Part of the inflorescence in the Gramineae. Spikes have two glumes
(bracts) with one or several flowers between them.

Spores Reproductive organs that detach from the parent plant and develop
into new plants.

Sub-spontaneous A term that appears in afforestation reports from the British mandate
to describe a naturalized alien species.

Syconium The flower and fruit of figs – an inflorescence in which the flowers
and fruit are enclosed in a jug-like succulent receptacle.

Vegetative Asexual reproduction via parts of the plant such as rhizomes, bulblets
propagation or or shoots, that detach from the parent plant and develop into new,
reproduction independent individuals.

HERBICIDES
2,4-D A systemic herbicide (=non-selective) licensed for use in Israel(1).
Affects native species(2)

Chlorsulfuron A systemic herbicide (=non-selective) licensed for use in Israel(1);

(trade name: Glean). Affects native species(2)

(1) From: The Ministry of Agriculture and Rural Development, Flora Protection and Inspection Services (2008) List of herbicides
approved for sale in Israel, January 2008, 80 pp.

(2) Is not permitted for use without prior approval from the Ministry of Agriculture and Rural Development and consultation with
the Israel Nature and Parks Authority.
Diquat A systemic herbicide (=non-selective) licensed for use in Israel(1)
(trade name: Reglone). Used to treat weeds in aquatic systems
(rivers, lakes). Affects native species(2)

Diuron A systemic herbicide (=non-selective) licensed for use in Israel(1)

(trade name: Diurex)

Fluridone A selective herbicide for aquatic systems whose use is forbidden in

Israel(1). Should not be applied in Israel.

Glyphosate Non-selective herbicide licensed for use in Israel(1) (trade name:

Roundup). A study in Israel showed that concentrated chemical
treatment of Acacia saligna with this herbicide is very effective in
neutralizing trees and seedlings. Affects native species(2)

Hexazinone Non-selective herbicide licensed for use in Israel(1) (trade name:

Velpar). Affects native species(2)

Paraquat A non-selective herbicide licensed for use in Israel(1). Affects native

species(2)

Picloram A selective herbicide licensed for use in Israel(1) (trade name: Tordon
181
101). Affects native species(2)

Triclopyr A selective herbicide licensed for use in Israel(1) (trade name: Garlon,
Crossbow, Grazon). Affects native species and can harm water
sources(2) .

ABBREVIATIONS

ISSG The Invasive Species Specialist Group is a global network of scientific

and policy experts on invasive species, organized under the auspices
of the Species Survival Commission (SSC) of the International Union
for Conservation of Nature (IUCN). The ISSG was founded in 1994.

IUCN International Union for Conservation of Nature. Founded in 1948 and

includes government and NGO members.

National A list published in 2000 by the Australian Government Department of

Environmental the Environment that includes 28 alien plant species in Australia that
(Alert) List are still in the early stages of invasion but have a high invasive
potential and are a significant threat to native ecosystems unless
managed as soon as possible.

WONS Weeds of National Significance – A list published in 1999 by the

Australian government. The list includes 21 species that have been
determined to present a serious threat to Australian ecosystems.

(1) From: The Ministry of Agriculture and Rural Development, Flora Protection and Inspection Services (2008) List of herbicides
approved for sale in Israel, January 2008, 80 pp.

(2) Is not permitted for use without prior approval from the Ministry of Agriculture and Rural Development and consultation with
the Israel Nature and Parks Authority.
182
References
1. AGENCE MEDITERRANEENNE de l’ENVIRONNEMENT (2003) Griffes de sorcière. Fiche No 7.
Ame-lr.org/publications/espaces/plantesenvahissantes/pdf/fiche07.pdf

2. AGUIAR F.C., FERREIRA M.T & MOREIRA I. (2001) Exotic and native vegetation establishment
following channelization of a western Iberian river. Regulated rivers: Research & Management
17:509-526.

3. AGUIAR F.C., FERREIRA M.T., ALBUQUERQUE A. & BERNEZ I. (2005) Invasibility Patterns of Knotgrass
(Paspalum distichum) in Portuguese Riparian Habitats. Weed Technology 19(3):509-516.

4. AHMED R., HOQUE A.T.M.R. & HOSSAIN M.K. (2008) Allelopathic effects of leaf litters of Eucalyptus
camaldulensis on some forest and agricultural crops. Journal of Forestry Research 19(1):19-24.

5. AJUONU O. & NEUENSCHWANDER P. (2003) Release, establishment, spread and impact of the
weevil Neohydronomus affinis (Coleoptera: Curculionidae) on water lettuce (Pistia stratiotes) in
Benin, West Africa. African Entomology 11(2):205–211.

6. AJUONU O., SCHADE V., VELTMAN B., SEDJRO K. & NEUENSCHWANDER P. (2003) Impact of the
weevils Neochetina eichhorniae and N. bruchi (Coleoptera: Curculionidae) on water hyacinth, 183
Eichhornia crassipes (Pontederiaceae), in Benin, West Africa. African Entomology 11(2):153–161.

7. AKMAN Y., KURT L., DEMIRYÜREK E, QUEZEL P., KURT F., EVREN H. & KÜÇÜKÖDÜK M. (1998) Les
groupements à Pinus brutia sur roches ultra-basiques et calcaires dans la région de Marmaris
et Bodrum (Mugla), à l’étage thermo-méditerranéen du sud-ouest anatolien (Turquie). Ecologia
Mediterranea 24(1):63-71.

8. AL-HUMAID A. I. & WARRAG M.O.A. (1998) Allelopathic effects of mesquite (Prosopis juliflora)
foliage on seed germination and seedling growth of bermudagrass (Cynodon dactylon). Journal of
Arid Environments 38(2):237-243.

9. ALBANO PÉREZ E., COETZEE J.A., RUIZ TÉLLEZ T., HILL M.P. (2011) A first report of water hyacinth
(Eichhornia crassipes) soil seed banks in South Africa. South African Journal of Botany, in press

10. ALMON N. (2005) Survey of the spread of Ambrosia sp. in the Alexander River basin. Presented to
the Director General of the Ministry of Agriculture, Mr.Yossi Yishai. Sharon Drainage Authority, January 2005, 6
pp. (In Hebrew)

11. ALSTON K.P. & RICHARDSON D.M. (2006) The roles of habitat features, disturbance, and distance
from putative source populations in structuring alien plant invasions at the urban/wildland
interface on the Cape Peninsula, South Africa. Biological Conservation 132(2):183-198.

12. ANGLISTER N., MOTRO U. & YOM-TOV Y. (2005) The effects of Acacia saligna and dune stabilization
on rodent populations in Ashdod-Nitzanim sands. Conference abstract of the world conference on
ecological retoration, Zaragoza, Spain 12-18 September 2005, p.27.

13. ANONYMOUS (2003) Karroo thorn (Acacia karroo). The Commonwealth Department of the
Environment, Water, Heritage and the Arts, and the CRC for Australian Weed Management. p.6.

14. ANONYMOUS (2003) Parkinsonia (Parkinsonia aculeata). The Commonwealth Department of the
Environment, Water, Heritage and the Arts, and the CRC for Australian Weed Management. p.8.

15. ANONYMOUS (2007a) Biological control of water lettuce, Pistia stratiotes. CSIRO (Australian
Commonwealth Scientific and Research Organization). http://www.csiro.au/resources/BiologicalControlWat
erLettuce.html [accessed 17 February, 2010].

16. ANONYMOUS (2007b) Ipomoea aquatica Forssk. http://www.texasinvasives.org/invasives_database/

detail.php?symbol=IPAQ [accessed 05 October, 2009].
 17. AREVALO J.M., DELGADO J.D., OTTO R., NARANJO A., SALAS M., FERNANDEZ-PALACIOS J.M. (2005)
Distribution of alien vs. native plant species in roadside communities along an altitudinal
gradient in Tenerife and Gran Canaria (Canary Islands). Perspectives in Plant Ecology, Evolution and
Systematics 7:185–202.

18. ARIANOUTSOU M., BAZOS I., DELIPETROU P. & KOKKORIS Y. (2010) The alien flora of Greece:
taxonomy, life traits and habitat preferences. Biological Invasions 12:3525-3549.

19. ARIANOUTSOU M., DELIPETROU P., CELESTI-GRAPOW L., BASNOU C., BAZOS I., KOKKORIS Y., BLASI
C. & VILA M. (2010) Comparing naturalized alien plants and recipient habitats across an east–west
gradient in the Mediterranean Basin. Journal of Biogeography 37:1811-1823.

20. ARIATI S.R., MURPHY D.J., GARDNER S. & LADIGES P.Y. (2007) Morphological and genetic variation
within the widespread species Acacia victoriae (Mimosaceae). Australian Systematic Botany 20:54-62.

21. BABIKER A. G. T., NAGAT E. M. & AHMED E. A. M. (2011) Mesquite (Prosopis juliflora): A threat to
agriculture and pastoralism in Sudan. In Brunel S, Uludag A, Fernandez-Galiano E, Brundu G (Eds.)
Proceedings of the 2nd International Workshop on Invasive Plants in the Mediterranean Type Regions of the
World, 2010-08-02/06, Trabzon, Turkey pp. 283-287.

22. BALLERO M., ARIU A., FALAGIANI P. & PIU G. (2003) Allergy to Ailanthus altissima (tree of heaven
pollen). Allergy 58(6):532-533.

23. BARRATT B.I.P., HOWARTH F.G., WITHERS T.M., KEAN J.M., RIDLEY G.S. (2010) Progress in risk
184 assessment for classical biological control. Biological Control 52:245-254.

24. BATCHER M. (2000) Element stewardship abstract for Melia azedarach. The Nature Conservancy’s
Wildland Invasive Species Program, edited by M.Tu & J.M. Randall. p.7. http://www.imapinvasives.org/GIST/ESA/
esapages/documnts/meliaze.pdf [accessed 08 September, 2009].

25. BERHANU A. & TESFAYE G. (2006) The Prosopis dilemma, impacts on dryland biodiversity and some
controlling methods. Journal of the Drylands 1(2):158-164.

26. BERNEZ I., FERREIRA M.T., ALBUQUERQUE A. & AGUIAR F. (2005) Relations between river plant
richness in the Portuguese floodplains and the widespread water knotgrass (Paspalum
paspaloides). Hydrobiologia 551:121-130.

27. BHAGWAT S.A., BREMAN E., THEKAEKARA T., THORNTON T.F. & WILLIS K.J. (2012) A Battle Lost?
Report on Two Centuries of Invasion and Management of Lantana camara L. in Australia, India
and South Africa. PLoS ONE 7(3) in press.

28. BLAMEY M. & GREY-WILSON C. (1993) Mediterranean wild flowers. Harper-Collins, p.560.

29. BLECHER M. (2005) Propagation of exotic trees by animals in the Dead Sea region and the Judean
Desert. Proceeding of the Conference on Biological Invasions in Israeli Ecosystems, pp. 28-29. (In Hebrew).

30. BLECHER M. & BLECHER A. (2005) Spread of exotic trees in oases in the Dead Sea region. Abstract
from Conference on Biological Invasions in Israeli Ecosystems, pp. 27-28. (In Hebrew)

31. BOLAND G.J. & BRIMNER T. (2004) Nontarget effects of biological control agents. New Phytologist
163(3):455-457.

32. BOSSARD C.C., RANDALL J.M. & HOSHOVSKY M.C. (2000) Invasive plants of California’s wildlands.
Univeristy of California Press, London. p.360.

33. BOSSARD C., BROOKS M., DITOMASO J., RANDALL J., ROYE C., SIGG J., STANTON A., WARNER P. (2006)
California Invasive Plant Inventory. Published by the California Invasive Plant Council, February 2006.
p.39.

34. BOTHMA A. (2002) Allelopathic potential of silverleaf nightshade (Solanum eleagnifolium Cav.).
MSc (Agric.) Horticulture. University of Pretoria, Pretoria (ZA).

35. BOYD J.W. & MURRAY D.S. (1982) Growth and development of silverleaf nightshade (Solanum
elaeagnifolium). Weed Science 30:238-243.
36. BRIESE D.T. (2004) Weed biological control: applying science to solve seemingly intractable
problems. Australian Journal of Entomology 43:304-317.

37. BRIMNER T.A. & BOLAND G.J. (2003) A review of the non-target effects of fungi used to biologically
control plant diseases. Agriculture, Ecosystems and Environment 100:3-16.

38. BURN A. (2003) The herbicide handbook: Guidance on the use of herbicides on nature
conservation sites. English Nature in association with FACT. Perterborough, UK.

39. CALEY P., GROVES R.H, BARKER R. (2007) Estimating the invasion success of introduced plants.
Diversity and Distributions 14(2):196-203.

40. CALVIÑO-CANCELA M., RUBIDO-BARÁ M.,VAN ETTEN E.J.B. (2012) Do eucalypt plantations provide
habitat for native forest biodiversity? Forest Ecology & Management 270:153-162.

41. CARSON W.P. HOVICK S.M.BAUMERT A.J. BUNKER D.E. & PENDERGAST T.H. (2008) Evaluating
the post-release efficacy of invasive plant biocontrol by insects: a comprehensive approach.
Arthropod-Plant Interactions 2:77-86.

42. CASTILLO J.M., LEIRA-DOCE P., CARRION-TACURI J., MUNOZ-GUACHO E., ARROYO-SOLIS A.,
CURADO G., DOBLAS D., RUBIO-CASAL A.E., ÁLVAREZ-LOPEZ A.A., REDONDO-GOMEZ S., BERJANO
R., GUERRERO G., DE CIRES A., FIGUEROA E. & TYE A. (2007) Contrasting strategies to cope with
drought by invasive and endemic species of Lantana in Galapagos. Biodiversity and Conservation
16(7):2123-2136.
185
43. CATFORD J.A., DOWNES B.J., GIPPEL C.J. & VESK P.A. (2011) Flow regulation reduces native plant
cover and facilitates exotic invasion in riparian wetlands. Journal of Applied Ecology 48(2):432-442.

44. CENTER for ECOLOGY & HYDROLOGY (2004) Information sheet 22: Azolla filiculoides,Water fern. Natural
Environment Research Council. Center for Aquatic Plant Management. http://www.nerc-wallingford.ac.uk/
research/capm/pdf%20files/22%20Azolla%20filiculoides.pdf [accessed 31 July, 2009].

45. CHALAK M., HEMERIK L.,VAN DER WERF W., RUIJS A. & VAN IERLAND E.C. (2010) On the risk of extinction
of a wild plant species through spillover of a biological control agent: Analysis of an ecosystem compartment
model. Ecological Modelling 221(16):1934-1943.

46. CHALAK M., RUIJS A. & VAN IERLAND E.C. (2011) Biological control of invasive plant species:
A stochastic analysis. Weed Biology & Management 11:137–151.

47. CHEN S.H., WENG S.H. & WU M.J. (2008) The Umbrella Sedge in Taiwan Taiwania 53(3):311-315.

48. CHENG W., XUEXIU C., HONGJUAN D., DIFU L. & JUNYAN L. (2008) Allelopathic inhibitory effect of
Myriophyllum aquaticum (Vell.) Verdc. On Microcystis aeruginosa and its physiological mechanism. Acta
Ecologica Sinica 28(6):2595-2603.

49. CHIKWENHERE G.P & FORNO I.W. (1991) Introduction of Neohydronomus affinis for Biological
Control of Pistia stratiotes in Zimbabwe. Journal of Aquatic Plant Management 29:53-55.

50. CHRISTEN D. & MATLACK G. (2006) The Role of Roadsides in Plant Invasions: a Demographic Approach.
Conservation Biology 20(2):385–391.

51. CHRISTEN D.C. & MATLACK G.R. (2009) The habitat and conduit functions of roads in the spread
of three invasive plant species. Biological Invasions 11:453-465.

52. COHEN O., RIOV J., KATAN J., GAMLIEL A. & BAR (KUTIEL) P. (2008) Reducing persistent seed banks of
invasive plants by soil solarization – the case of Acacia saligna. Weed Science 56(6):860-865.

53. COHEN O., RIOV J. & BAR P. (2010a) Rehabilitation of the Einot Gibton wetland after biological
invasion by Acacia saligna. Ecology & Environment 1:8-9. (in Hebrew)

54. COHEN O., RON M., MELIHI Y., LAZAR A., UZAN A., VALCHAK M., KESHET N. & RIOV J. (2010b)
Rehabilitation of natural vegetation in the Einot Gibton wetland Nature Reserve following
invasion by Acacia saligna. Proceedings of the 51st annual conference of the Israeli geography association,
the Hebrew University, 5-7 December 2010, pp 158-159. (in Hebrew).
 55. CONSER C. & CONNOR E.F. (2009) Assessing the residual effects of Carpobrotus edulis invasion,
implications for restoration. Biological Invasions 11:349-358.

56. CONSTÁN-NAVA S., BONET A., PASTOR E., LLED M.J. (2010) Long-term control of the invasive tree
Ailanthus altissima: Insights from Mediterranean protected forests. Forest Ecology & Management
260(6):1058-1064.

57. CRONK Q. C. B. & FULLER J. L. (2001) Plant Invaders: the Threat to Natural Ecosystems. Earthscan
Publications, London, UK. p.241.

58. CSIRO (2004) Eucalyptus camaldulensis Dehnh. http://www.anbg.gov.au/cpbr/WfHC/Eucalyptus-

camaldulensis/ [accessed 06 September, 2009]

59. D’ANTONIO C.M. (1993) Mechanisms controlling invasion of coastal plant communities by the
alien succulent Carpobrotus edulis. Ecology 74(1):83-95.

60. DAEHLER C.C., DENSLOW J.S., ANSARI S. & KUO H.C. (2004) A Risk-Assessment System for
Screening Out Invasive Pest Plants from Hawaii and Other Pacific Islands. Conservation Biology
18(2):360-368.

61. DAFNI A. & HELLER D. (1990) Invasions of adventive plants in Israel. In: Di Castri F., Hansen A.J. &
Debussche M. (Eds.) Biological invasions in Europe and the Mediterranean basin. Kluwer, p.463. pp.135-160.

62. DAISIE (2008) Handbook of Alien Species in Europe (Invading Nature – Spinger Series in Invasion
186 Ecology). P. 400.

63. DANIN A. (1976) Notes on four adventives composites in Israel. Notes from the Royal Botanic Garden
Edinburgh 34(3):403-410.

64. DANIN A. (1994) Contribution to the flora of Israel. VI Stipagrostis drarii and Ambrosia
confertiflora and tenuifolia, new records from Israel. Israel Journal of Vegetation Sciences 42:59-61.

65. DANIN A. (2004) Distribution Atlas of Plants in the Flora Palaestina Area. The Israel Academy of
Sciences and Humanities, Jerusalem, Israel. p.517.

66. DANIN A. & DUFOUR-DROR J.M. (2008) Cyperus odoratus L. In: Greuter W. & Raus T. (Eds) Med-checklist
Notulae 27, Willdenowia 38, p.469.

67. DAUER J.T., MORTENSEN D.A., VANGESSEL M.J. (2007) Temporal and spatial dynamics of long-
distance Conyza canadensis seed dispersal. Journal of Applied Ecology 44(1):105-114.

68. DAUER J.T., MORTENSEN D.A., LUSCHEI E.C, ISARD S.A., SHIELDS E., VAN-GESSEL M.J. (2009) Conyza
canadensis seed ascent in the lower atmosphere. Agricultural & Forest Meteorology 149(3-4):526-534.

69. DE CLERCQ P., MASON P.G. & BABENDREIER D. (2011) Benefits and risks of exotic biological control
agents. BioControl 56:681-698.

70. DEAN W.R.J., ANDERSON M.D., MILTON S.J. & ANDERSON T.A. (2002) Avian assemblages in native
Acacia and alien Prosopis drainage line woodland in the Kalahari, South Africa Journal of Arid
Environments 51:1–19.

71. DEGEN A.A., BECKER K., MAKKER H.P.S. & BOROWY N. (1995) Acacia saligna as a fodder tree for
desert livestock and the interaction of its tannins with fibre fractions. Journal of the Science of Food
and Agriculture 68(1): 65-71.

72. DEGEN A.A., BLANKE A., BECKER K., KAM M., BENJAMIN R.W. & MAKKER H.P.S (1997) The nutritive
value of Acacia saligna and Acacia salicina for goats and sheep. Animal Science Pencaitland 64(2):
253-259.

73. DEN HOLLANDER N.G., SCHENK I.W., DIOUF S., KROPFF M.J. & PIETERSE A.H. (1999) Survival strategy
of Pistia stratiotes L. in the Djoudj National Park in Senegal. Hydrobiologia 415:21-27.

74. DI CASTRI F., GROVES R.H., KRUGER F.J., REJMÁNEK M., WILLIAMSON M., DRAKE J.A. & MOONEY H.A.
(1989) Biological invasions: A global perspective. Scope 37. John Wiley & Sons. p. 550.
75. DJURDJEVIC´ L., MITROVIC´ M., GAJIC´ G., JARIC´ S., KOSTIC´ O., OBERAN L., PAVLOVIC´ P. (2011)
An allelopathic investigation of the domination of the introduced invasive Conyza canadensis L.
Flora 206(11):921-927.

76. DONALD D.G.M. & KIRBY-SMITH J.H. (1983) Chemical control of phyllode Acacias in the
SW Cape. In: Neser S. & Cairns A.L.P (Eds) Proceedings of the fifth national weeds conference of South-
Africa, Stellenbosch.

77. DONNELLY G.P (2000) Biology and host specificity of Rhinacloa callicrates Herring (Hemiptera:
Miridae) and its introduction and establishment as a biological control agent of Parkinsonia
aculeata L. (Caesalpiniaceae) in Australia. Australian Journal of Entomology 39(2):89-94.

78. DONNELLY M.J., GREEN D.M. & WLTERS L.J. (2008) Allelopathic effects of fruits of the Brazilian
pepper Schinus terebinthifolius on growth, leaf production and biomass of seedlings of the red
mangrove Rhizophora mangle and the black mangrove Avicennia germinans. Journal of Experimental
Marine Biology and Ecology 357:149-156.

79. DRAY F.A. & CENTER T.D. (1989) Seed production by Pistia stratiotes L. (Water Lettuce) in the
United States. Aquatic Botany 33:155-160.

80. DU PUY D.J., LABAT J.N., RABEVOHITRA R.,VILLIERS J.F., BOSSER J. & MOAT J. (2002) The Leguminosae
of Madagascar. Royal Botanic Gardens, Kew.

81. DUFOUR-DROR J.M. & DANIN A. (2004) Acacia paradoxa DC. In: Greuter W. & Raus T. (Eds) Med-checklist 187
Notulae, 22. Willdenowia 34:71-80, p.75.

82. DUFOUR-DROR J.M. & ERTAS A. (2004) Bioclimatic perspectives in the distribution of Quercus
ithaburensis Decne. Subspecies in Turkey and in the Levant. Journal of Biogeography 31(3):461-474.

83. DUFOUR-DROR J.M. (2007a) Reducing the invasion process of Acacia saligna in natural areas with
the chemical targeted control method. Final Report submitted to the Science & Conservation Division
of the Israel Nature and Parks Authority, 35 pp. (In Hebrew, summary in English).

84. DUFOUR-DROR J.M. (2007b) Reducing the proliferation of invasive plants in open and protected
spaces in the Mediterranean region in Israel. Final Report submitted to the Israel Ministry of
Environmental Protection, 97 pp. (In Hebrew).

85. DUFOUR-DROR J.M. (2009) Invasive plant species in Israel’s natural areas: Distribution, stages
of invasion, degrees of ecological threat and a tool for prioritizing control of invasive plant
populations. The Jerusalem Institute for Israel Studies – The Environmental Policy Center, the Israel Nature
and Parks Authority – Science and Conservation Division. The Jerusalem Institute, Publication no. 33. 108 pp.
(In Hebrew)

86. DUFOUR-DROR J.M. (2010) The control of Ambrosia confertiflora along the Alexander River
banks: An attempt to develop a control method based on the restoration of riparian natural
vegetation. Proceedings of the 38th annual conference of the Israel Society of Ecology and Environmental
Sciences, 20-22 June 2010, Ben-Gurion University.

87. DUFOUR-DROR J.M., FRAGMAN-SAPIR O., KAGAN S., YAACOBY T. & WALCZAK M. (2012) The least
wanted alien plant species in Israel. The Israel Ministry of Environment, in press. (In Hebrew).

88. ECONOMOU G., TZAKOU O., GANI A., YANNITSAROS A. & BILALIS D. (2002) Allelopathic effect of
Conyza albida on Avena sativa and Spirodela polyrhiza. Journal of Agronomy and Crop Science 188(4):
248-253.

89. EIJZENGA H. (2011) Vegetation change following rabbit eradication on Lehua Island, Hawaiian
Islands. In:Veitch, C. R.; Clout, M. N. and Towns, D. R. (eds.) Island invasives: eradication and management. IUCN,
Gland, Switzerland, 290-294.

90. EL-KEBLAWY A. & AL-RAWAI A. (2007) Impacts of the invasive exotic Prosopis juliflora (Sw.) D.C.
on the native flora and soils of the UAE. Plant Ecology 190(1):23-35.
 91. ELTON C.S. (1958) The Ecology of invasions by animals and plants. Methuen, London, UK.

92. ESSA S., DOHAI B. & KSIKSI T. (2006) Mapping dynamics of invasive Prosopis juliflora in the northern
emirates of the uae: an application of remote sensing and GIS. ISPRS Commission VII Mid-term
Symposium “Remote Sensing: From Pixels to Processes”, Enschede, the Netherlands, 8-11 May 2006.

93. EPPO (2005) EPPO data sheet on Invasive Plants – Myriophyllum aquaticum. www.eppo.org/QUARANTINE/
Pest_Risk_Analysis/PRAdocs-plants/draftds/05-11833%20DS%20Myriophyllum%20aquaticum.doc [accessed
16 August, 2009]

94. EPPO (2007) Solanum elaeagnifolium. Data sheets on quarantine pests. EPPO Bulletin 37:236-245.

95. EPPO (2007) Salvinia molesta (Salviniaceae), Giant Salvinia. www.eppo.org/QUARANTINE/Alert_List/

invasive_plants/Salvinia_molesta.htm [accessed 30 July, 2009].

96. EPPO (2008) Verbesina encelioides (Asteraceae). www.eppo.org/quarantine/Alert_List/invasive_plants/

Verbesina_encelioides.htm [accessed May 6th 2009].

97. ESLER K.J., HOLMES P.M., RICHARDSON D.M. & WITKOWSKI E.T.F. (2008) Riparian vegetation
management in landscapes invaded by alien plants: Insights from South Africa. South African Journal
of Botany 74(3):397-400.

98. ESSA S., DOHAI B. & KSIKSI T. (2006) Mapping dynamics of invasive Prosopis juliflora in the northern
emirates of the UAE: An application of remote sensing and GIS. ISPRS Commission VII Mid-term
188 Symposium “Remote Sensing: From Pixels to Processes”, Enschede, the Netherlands, 8-11 May 2006.

99. EVANS D.O. (2001) Sesbania sesban: Widely distributed multipurpose NFT. In: Roshetko J.M. (ed.)
Agroforestry Species and Technologies, pp. 157-158. Winrock International, Morilton, USA.

100. FAHN A., HELLER D. & AVISHAI M. (1998) The cultivated plants of Israel. Kibbutz Meuhad. 703 pp.
(In Hebrew)

101. FAITHFULL I. & BLOOD K. (2004) Karoo thorn and giraffe thorn: state prohibited weeds. Landcare
Notes. State of Victoria, Department of Sustainability and Environment. p. 4.

102. FAO (?) Pennisetum clandestinum Hochst. Ex Chiov. http://www.fao.org/ag/AGP/AGPC/doc/Gbase/

DATA/Pf000298.htm [accessed 20 July, 2009].

103. FEENSTRA K.R. & CLEMENTS D.R. (2008) Biology and Impacts of Pacific Island Invasive Species.
4. Verbesina encelioides, Golden Crownbeard (Magnoliopsida: Asteraceae). Pacific Science 62(2):
161-176.

104. FEINBURN-DOTHAN N. & DANIN A. (1991) Analytical flora of Eretz-Israel. CANA Publishing House
Ltd. 1040 pp. (In Hebrew)

105. FENSHAM R.J., FAIRFAX, R.J. & CANNELL R.J. (1994) The invasion of Lantana camara L. in Forty Mile
Scrub National Park, north Queensland. Australian Journal of Ecology 19(3):297-305.

106. FERNÁNDEZ-ZAMUDIOA R., GARCÍA-MURILLOA P., CIRUJANO S. (2010) Germination characteristics

and sporeling success of Azolla filiculoides Lamarck, an aquatic invasive fern, in a Mediterranean
temporary wetland. Aquatic Botany 93:89-92.

107. FERREIRO D., OROZCO J. P., MIRÓN C., REAL T., HERNÁNDEZ-MORENO D., SOLER F. & PÉREZ-LÓPEZ
M. (2010) Chinaberry Tree (Melia azedarach) Poisoning in Dog: A Case Report. Topics in Companion
Animal Medicine 25(1):64-67.

108. FERRITER A. (1997) Brazilian Pepper Management Plan for Florida. A report from the Florida Exotic
Pest Plant Council’s Brazilian Pepper Task Force. p. 26

109. FINNOFF D., SHOGREN J.F., LEUNG B. & LODGE D. (2006) Take a risk: Preferring prevention over
control of biological invaders. Ecological Economics 62(2):216-222.

110. FLORENTINE S.K., WESTBROOKE M.E.(2005) Invasion of the noxious weed Nicotiana glauca R.
Graham after an episodic flooding event in the arid zone of Australia. Journal of Arid Environments
60(4):531-545.
111. FLORENTINE S.K., WESTBROOKE M.E., GOSNEY K., AMBROSE G. & O’KEEFE M. (2006) The arid
land invasive weed Nicotiana glauca R. Graham (Solanaceae): Population and soil seed bank
dynamics, seed germination patterns and seedling response to flood and drought. Journal of Arid
Environments 66:218-230.

112. FLORIDA DEPARTMENT of ENVIRONMENTAL PROTECTION WEED ALERT (?) Water-spinach

(Ipomoea aquatica). Bureau of Invasive Plant Management. p.2. http://www.dep.state.fl.us/lands/invaspec/
3rdlevpgs/Wtrspnch.pdf [accessed 05 October, 2009].

113. FLORY S.L. & CLAY K. (2009) Effects of roads and forest successional age on experimental plant
invasions. Biological Conservation 142:2531-2537.

114. FOX J.E.D. (1995) A review of the ecological characteristics of Acacia saligna (Labill.) H. Wendl.
Mulga Research Center Journal 12:39-56.

115. FRAGMAN O., PLITMANN U., HELLER D. & SHMIDA A. (1999) Checklist and ecological data-base of
the flora of Israel and its surroundings. Israel Nature and Natural Parks protection Authority, Jerusalem.
107 pp.

116. FRAGMAN-SAPIR O. (2008) Flowers of Ein Gedi and the Dead Sea Coast – Plant Guide. Israel Nature
& Parks Authority. 225 pp. (In Hebrew)

117. GAERTNER M., DEN BREEYEN A., HUI C. & RICHARDSON D.M. (2009) Impacts of alien plant invasions
on species richness in Mediterranean-type ecosystems: a meta-analysis. Progress in Physical Geography
33(3):1–20.
189

118. GALIL J. & EISIKOITCH D. (1968) On the pollination ecology of Ficus religiosa in Israel. Phytomorphology
18:356-363.

119. GALON Y., DRORI Y., AVISHAI M., YAFFE Y. & HEL-OR Y. (1993) List of the ornamental plants in
Israel. Ministry of Agriculture Agriculture, Department of Plant Engineering, Gardening and Landscaping.
(In Hebrew).

120. GASSMANN A., COCK M.J.W., SHAW R. & EVANS H.C. (2006) The potential for biological control of
invasive alien aquatic weeds in Europe: a review. Hydrobiologia 570(1):217-222.

121. GEERTS S. & PAUW A. (2009) African sunbirds hover to pollinate an invasive hummingbird-
pollinated plant. Oikos 118(4):573-579.

122. GELBARD J.L. & BELNAP J. (2003) Roads as Conduits for Exotic Plant Invasions in a Semiarid
Landscape. Conservation Biology 17(2): 420–432.

123. GEORGE N., BYRNE M., MASLIN B. & YAN G. (2006) Genetic differentiation among morphological
variants of Acacia saligna (Mimosaceae). Tree Genetics & Genomes 2(2):109-119.

124. GERBER E., SCHAFFNER U., GASSMANN A., HINZ H.L., SEIER M. & MÜLLER-SCHÄRER H (2011)
Prospects for biological control of Ambrosia artemisiifolia in Europe: learning from the past Weed Research
51(6):559-573.

125. GIMENO I., VILA M., HULME P.E. (2006) Are islands more susceptible to plant invasion than
continents? A test using Oxalis pes-caprae L. in the western Mediterranean. Journal of Biogeography
33(9):1559-1565.

126. GLOBAL INVASIVE SPECIES DATABASE (2005) Pennisetum clandestinum. http://www.issg.org/database/

species/ecology.asp?si=183&fr=1&sts=&lang=EN [accessed 20 July, 2009].

127. GLOBAL INVASIVE SPECIES DATABASE (2005) Myriophyllum aquaticum (aquatic plant). www.issg.org/
database/species/ecology.asp?si=401&Fr=1&sts=&lang=EN [accessed 16 August, 2009].

128. GLOBAL INVASIVE SPECIES DATABASE (2006) Ipomoea aquatica. http://www.issg.org/database/species/

ecology.asp?fr=1&si=477 [accessed 05 October, 2009].

129. GLOBAL INVASIVE SPECIES DATABASE (2006) Melia azedarach. http://www.issg.org/database/species/

ecology.asp?si=636&fr=1&sts= [accessed 08 September, 2009].
 130. GLOBAL INVASIVE SPECIES DATABASE (2009) Nicotiana glauca (shrub). http://www.issg.org/database/
species/ecology.asp?si=1453&fr=1&sts=&lang=EN [accessed 20 July, 2009].

131. GOMEZ-APARICIO L., CANHAM C.D. (2008) Neighbourhood analyses of the allelopathic effects of
the invasive tree Ailanthus altissima in temperate forests Journal of Ecology 96 (3) , 447–458.

132. GOODEN B., FRENCH K., TURNER P.J. (2008) Invasion and management of a woody plant, Lantana
camara L., alters vegetation diversity within wet sclerophyll forest in southeastern Australia.
Forest Ecology and Management 257(3):960-967.

133. GOODEN B., FRENCH K., TURNER P.J. & DOWNEY P.O. (2009) Impact threshold for an alien plant
invader, Lantana camara L., on native plant communities. Biological Conservation 142(11):2631-2641.

134. GRATWICKE B. & MARSHALL B.E. (2001) The impact of Azolla filiculoides Lam. on animal
biodiversity in streams in Zimbabwe. African Journal of Ecology 39(2):216-218.

135. GROENTEMAN R., FOWLER S.V., SULLIVAN J.J. (2011) St. John’s wort beetles would not have been
introduced to New Zealand now: A retrospective host range test of New Zealand’s most
successful weed biocontrol agents. Biological Control 57:50-58.

136. GROVES R.H. (2006) Are some weeds sleeping? Some concepts and reasons. Euphytica 148:111-120.

137. GROVES R.H., PANETTA F.D. & VIRTUE J.G. (2001) Weed Risk Assessment. CSIRO Publishing, p.244.

138. GROVES R.H.C., HOSKING J.R., BATIANOFF G.N., COOKE D.A., COWIE I.D., JOHNSON R.W., KEIGHERY
190 G.J., LEPSCHI B.J., MITCHELL A.A., MOERKERK M., RANDALL R.P., ROZEFELDS A.C., WALSH N.G. &
WATERHOUSE B.M. (2003) Weed categories for natural and Agricultural ecosystem management. Canberra:
Bureau of Rural Sciences, p.194.

139. GROVES R.H., BODEN R. & LONSDALE M. (2005) Jumping the garden fence: Invasive garden plants
in Australia and their environmental and agricultural impacts. A CSIRO report for WWF-Australia.
p.173.

140. GUILLERM J.L., LE FLOC’H E., MAILLET J. & BOULET C. (1990) The invading weeds within the Western
Mediterranean. In: Di Castri F., Hansen A.J. & Debussche M. (Eds.) Biological invasions in Europe and the
Mediterranean basin. Kluwer, p.463. pp.61-84.

141. GUTTERIDGE R.C. (1994) The perennial Sesbania species. In: Gutteridge R.C. & Shelton H.M. (eds.)
Forage Tree Legumes in Tropical Agriculture. Pp.49-64, Tropical Grassland Society of Australia, Brisbane,
Australia.

142. HADJIKYRIAKOU G. & HADJISTERKOTIS E. (2002) The adventive plants of Cyprus with new records
of invasive species Zeitschrift Fur Jagdwissenschaft 48:59-71.

143. HAIGEN X., SHENG Q., ZHENGMIN H., JIANYING G., ZONGGUO H., HONGYING S., SHUNPING H.,
HUI D., HAIRONG W. & FANGHAO W. (2006) The status and causes of alien species invasion in China.
Biodiversity and Conservation 15(9):2893-2904.

144. HALVORSON W.L. & GUERTIN P. (2003) Factsheet for Heterotheca subaxillaris (Lam.) Britt.
& Rusby. US Geological Survey, ‘Weeds in the West’ project, Status of Introduced Plants in Southern Arizona
Parks. Southwest Biological Science Center, Sonoran Desert Field Station, University of Arizona, p. 25.

145. HAR-EDOM O.L. & STERNBERG M. (2009) Invasive species and climate change: Conyza canadensis
(L.) Cronquist as a tool for assessing the invasibility of natural plant communities along an
aridity gradient. Biological Invasions 12(7):1953-1960.

146. HARDWOOD E. & SYTSMA M. (2003) Risk Assessment for Cinese Water Spinach (Ipomoea
aquatica) in Oregon. Center for Lakes and Reservoirs, Portland State University. http://oregon.gov/OISC/
docs/pdf/ipaq_ra.pdf [accessed 05 October, 2009].

147. HARLEY K.L.S., KASSULKE R.C., SANDS D.P.A. & DAY M.D. (1990) Biological control of water lettuce,
Pistia stratiotes [Araceae] by Neohydronomus affinis [Coleoptera: Curculionidae]. BioControl
35(3):363-374.
148. HARRINGTON M.G. & GADEK P.A. (2009) A species well travelled – the Dodonaea viscosa
(Sapindaceae) complex based on phylogenetic analyses of nuclear ribosomal ITS and ETSf
sequences. Journal of Biogeography 36:2313-2323.

149. HENDERSON L. (2001) Alien Weeds and Invasive Plants: A complete guide to declared weeds and
invaders in South Africa. Plant Protection Research Institute Handbook No.12, ARC LNR, p. 300.

150. HEYLIGERS P.C. (1998) Some New South Wales coastal plant distributions: a comparison of
herbarium records with transect survey data. Cunninghamia 5(3):645-664.

151. HILL M.P. & CILLIERS C.J. (1999) Azolla filiculoides Lamarck (Pteridophyta: Azollaceae), its status in
South Africa and control. Hydrobiologia 415:203-206.

152. HOLM L., DOLL J., HOLM E., PANCHO J. & HERBERGER J. (1997) World weeds: Natural histories and
distribution. Wiley, 1152 p.

153. HOLMES P.M. (1989) Decay rates in buried alien Acacia seed populations of different density.
South African Journal of Botany 55(3):299-303.

154. HOLMES P.M. (2001) Shrubland restoration following woody alien invasion and mining: Effects of
topsoil depth, seed source and fertilizer addition. Restoration Ecology 9(1):71-84.

155. HOLMES P.M., MACDONALD I.A.W. & JURITZ J. (1987). Effects of clearing treatment on seed banks
of the alien invasive shrubs Acacia saligna and Acacia cyclops in the southern and south-western
191
Cape, South Africa. Journal of Applied Ecology 24(3): 1045-1052.

156. HOLMES P.M. & MOLL E.J. (1990) Effect of depth and duration of burial on alien Acacia saligna and
Acacia cyclops seeds. South African Journal of Ecology 1:12-17.

157. HOLMES P.M., ESLER K.J., RICHARDSON D.M. & WITKOWSKI E.T.F. (2008) Guidelines for improved
management of riparian zones invaded by alien plants in South Africa. South African Journal of Botany
74(3):538-552.

158. HOLMQUIST J.G., SCHMIDT-GENGENBACH J. & SLATON M.R. (2011) Influence of invasive palms on
terrestrial arthropod assemblages in desert spring habitat. Biological Conservation 144:518-525.

159. HOWARD V. (2009) Pistia stratiotes. US Geological Survey Nonindigenous Aquatic Species Database,
Gainesvill, FL. http://nas.er.usgs.gov/queries/FactSheet.asp?speciesID=1099 [accessed 17 February, 2010].

160. HSIAO A.I. & HUANG W.Z. (1989) Effects of flooding on rooting and sprouting of isolated stem
segments and on plant growth of Paspalum distichum L. Weed Research 29(5):335-344.

161. HULME P.E. (2006) Beyond control: wider implications for the management of biological invasions.
Journal of Applied Ecology 43(5):835-847.

162. HUSSEY B.M.J., KEIGHERY G.J., DODD J., LLOYD S.G. & COUSENS R.D. (2007) Western weeds:
A guide to the weeds of Western Australia. 2nd edition. The Weed Society of Western Australia, p. 294.

163. HUSSNER A. (2006) NOBANIS – Invasive Alien Species Fact Sheet – Azolla filiculoides. – From: Online
Database of the North European and Baltic Network on Invasive Alien Species – NOBANIS www.nobanis.org
[accessed 02 August, 2009].

164. IMPSON F.A.C., MORAN V.C. & HOFFMANN J.H. (2004) Biological control of an alien tree, Acacia
cyclops, in South Africa: impact and dispersal of a seed-feeding weevil, Melanterius servulus.
Biological Control 29:375-381.

165. IMPSON F.A.C., KLEINJAN C.A., HOFFMANN J.H., POST J.A. & WOOD A.R. (2011) Biological control of
Australian Acacia species and Paraserianthes lophantha (Willd.) Nielsen (Mimosaceae), in South
Africa. Afr Entomol 19(2):186–207.

166. INDERJIT, CADOTTE M.W, COLAUTTI R.I. (2005) The Ecology of biological invasions: past, present
and future. pp. 19-43. In Inderjit (ed.) Invasive plants: Ecological and agricultural aspects. P. 283, Birkhäuser.
 167. IPONGA D.M., MILTON S.J. & RICHARDSON D.M. (2008) Superiority in competition for light:
A crucial attribute defining the impact of the invasive alien tree Schinus molle (Anacardiaceae)
in South African savanna. Journal of Arid Environments 72(5):612-623.

168. IPONGA D.M., MILTON S.J. & RICHARDSON D.M. (2009) Soil type, microsite, and herbivory influence
growth and survival of Schinus molle (Peruvian pepper tree) invading semi-arid African savanna.
Biological Invasions 11:159-169.

169. JADHAV A., HILL M. & BYRNE M. (2008) Identification of a retardant dose of glyphosate with
potential for integrated control of water hyacinth, Eichhornia crassipes (Mart.) Solms-Laubach.
Biological Control 47:154:158.

170. JENSEN W.I. & ALLEN J.P. (1981) Naturally occurring and experimentally induced castor bean
(Ricinus communis) poisoning in ducks. Avian Disease 25(1):184-194.

171. KAIRO M., ALI B., CHEESMAN O., HAYSOM K. & MURPHY S. (2003) Invasive Species in the Caribbean
Region. Report to the Nature Conservancy, CAB International. p.134.

172. KALWIJ J.M., MILTON S.J. & MCGEOCH M.A. (2008) Road verges as invasion corridors? A spatial
hierarchical test in an arid ecosystem. Landscape Ecology 23(4):439-451.

173. KARLSSON L.M. & MILBERG P. (2007) Comparing after-ripening response and germination
requirements of Conyza canadensis and C. bonariensis (Asteraceae) through logistic functions.
192 Weed Research 47(5):433-441.

174. KEANE M. & CRAWLEY M.J. (2002) Exotic plant invasions and the enemy release hypothesis. Trends
in Ecology & Evolution 17(4):164-170.

175. KELLY, M., B. PETERSON, et al. (2007). Drilling and injecting two invasive palm species. California
Invasive Plant Council Symposium 2007. San Diego, CA, California Invasive Plant Council.

176. KEW ROYAL BOTANICAL GARDEN http://www.kew.org/plants-fungi/Dodonaea-viscosa.htm

177. KIM Y.O., JOHNSON J.D. & LEE E.J (2007) Phytotoxicity of Phytolacca americana Leaf Extracts on
the Growth, and Physiological Response of Cassia mimosoides. Journal of Chemical Ecology 31(12)
:2963-2974.

178. KONG C.H., WANG P., ZHANG C.X., ZHANG M.X. & HU F. (2006) Herbicidal potential of
allelochemicals from Lantana camara against Eichhornia crassipes and the alga Microcystis
aeruginosa. Weed Research 46, 290–295.

179. KOTOULA-SYKA E. (2011) Solanum elaeagnifolium, an increasing problem in Greece. In Brunel S,

Uludag A, Fernandez-Galiano E, Brundu G (Eds.) Proceedings of the 2nd International Workshop on Invasive
Plants in the Mediterranean Type Regions of the World, 2010-08-02/06, Trabzon, Turkey pp. 400-403.

180. KOWARIK I. & SAUMEL I. (2007) Biological flora of Central Europe: Ailanthus altissima (Mill.)
Swingle. Perspectives in Plant Ecology Evolution and Systematics 8(4):207-237.

181. KOWARIK I. & SAUMEL I. (2008) Water dispersal as an additional pathway to invasions by the
primarily wind-dispersed tree Ailanthus altissima. Plant Ecology 198:241-252.

182. KULL C.A., SHACKLETON C.M., CUNNINGHAM P.S., DUCATILLON C., DUFOUR-DROR J.M., ESLER K.J.,
FRIDAY J.B., GOUVEIA A.C., GRIFFIN A.R., MARCHANTE E.M., MIDGLEY S.J., PAUCHARD A., RANGAN
H., RICHARDSON D.M., RINAUDO T., TASSIN J., URGENSON L.S., VON MALTITZ G.P., ZENNI R.D.,
ZYLSTRA M.J. (2011) Adoption, use, and perception of Australian acacias around the world. Diversity
& Distribution 17:822-836.

183. LAMBDON P. (2006) Oxalis pes-caprae. Delivering Alien Invasive Species Inventories for Europe (DAISIE).
p.3. http://www.montsevila.org/Species/Oxalis_pes-caprae.pdf [accessed 03 August, 2009].

184. LAND PROTECTION (2007) Water lettuce, Pistia stratiotes. Factsheet, Biosecurity, The State of
Quennsland (Department of Primary Industries and Fisheries). 4 p.
185. LAWRENCE J.G., COLWELL A. & SEXTON O.J. (1991) The ecological impact of a allelopathy in
Ailanthus altissima (Simaroubaceae). American Journal of Botany 78(7):948-958.

186. LE MAITRE D.C., GAERTNER M., MARCHANTE E., ENS E.J., HOLMES P.M., PAUCHARD A., O’FARRELL
P.J., ROGERS A.M., BLANCHARD R., BLIGNAUT J. & RICHARDSON D.M. (2011) Impacts of invasive
Australian acacias: implications for management and restoration. Diversity & Distribution
17:1015-1029.
187. LE ROUX J.J., BROWN G.K., BYRNE M., NDLOVU J., RICHARDSON D.M., THOMPSON G.D & WILSON
J.R.U. (2011) Phylogeographic consequences of different introduction histories of invasive
Australian Acacia species and Paraserianthes lophantha (Fabaceae) in South Africa. Diversity &
Distribution 17:861-871.

188. LIFSHITZ N. & BIGER G. (2004) The current distribution of the Turkish pine (Pinus brutia):
A renewed evaluation in the light of dendroarchaeological and dendrohistorical evidence. Forest
5-6:19-24. (In Hebrew).
189. LIN H.F., ALPERT P. & YU F.H. (2012) Effects of fragment size and water depth on performance of
stem fragments of 6 the invasive, amphibious, clonal plant Ipomoea aquatica. Aquatic Botany, in
press.
190. LISANEWORK N. & MICHELSEN A. (1993) Allelopathy in agroforestry systems: the effects of leaf
extracts of Cupressus lusitanica and three Eucalyptus spp. on four Ethiopian crops. Agroforestry
Systems 21(1):63-74. 193
191. LONSDALE W.M. (1999) Global patterns of plant invasions and the concept of invasibility. Ecology
80(5):1522-1535.
192. LOWE S., BROWNE M., BOUDJELAS S. & De POORTER M. (2004) 100 of the world’s worst invasive
alien species. A selection from the global invasive species database. 2nd edn.The invasive species speciality group
(ISSG) a specialist group of the species survival commission (SSC) of the world conservation union (IUCN).
http://www.issg.org/pdf/publications/worst_100/english_100_worst.pdf
193. MALCOLM G.M., BUSH D.S. & RICE S.K. (2008) Soil nitrogen conditions approach preinvasion
levels following restoration of nitrogen-fixing Black Locust (Robinia pseudoacacia) stands in a
Pine–Oak ecosystem. Restoration Ecology 16(1):70-78.
194. MALTEZ-MOURO S., MAESTRE F.T. & FREITAS H. (2010) Weak effects of the exotic invasive
Carpobrotus edulis on the structure and composition of Portuguese sand-dune communities.
Biological Invasions 12(7):2117-2130.

195. MARCHANTE E., KJØLLER A., STRUWE S. & FREITAS H. (2008) Short- and long-term impacts of Acacia
longifolia invasion on the belowground processes of a Mediterranean coastal dune ecosystem.
Applied Soil Ecology 40:210-217.

196. MARCO A. LAVERGNE S. DUTOIT T. & BERTAUDIERE-MONTES V. (2010) From the backyard to
the backcountry: how ecological and biological traits explain the escape of garden plants into
Mediterranean old fields. Biological Invasions 12:761-779.
197. MARINGER J., WOHLGEMUTH T., NEFF C., PEZZATTI G.B., CONEDERA M. (2012) Post-fire spread of
alien plant species in a mixed broad-leaved forest of the Insubric region. Flora 207(1):19-29.
198. MARTINS V.F., GUIMARÃES P.R., HADDAD C.R.B. SEMIR J. (2009) The effect of ants on the seed
dispersal cycle of the typical myrmecochorous Ricinus communis. Plant Ecology 205(2):213-222.
199. MASAKA K. & YAMADA K. (2009) Variation in germination character of Robinia pseudoacacia L.
(Leguminosae) seeds at individual tree level. Journal of Forest Research 14:167-177.
200. MASLIN B. R. (2001) Wattle – Acacias of Australia. Department of Conservation and Land Management
(CALM), Perth, WA, CSIRO Publishing, Collingwood,VIC.
201. MASLIN B.R. & McDONALD M.W. (2003) Acacia victoriae. In B.R. Maslin & M.W. McDonald (eds) Acacia
search – Evaluation of Acacia as a woody crop option for southern Australia. pp.222-226. Joint Venture Agroforestry
program. RIRDC Publication No 03/017, Project No CAL-7A.
 202. MASLIN B. R. & McDONALD M.W. (2004a) Acacia saligna (Labill.) H. Wendl. In Maslin B.R. & McDonald
M.W. (Eds) AcaciaSearch - Evaluation of Acacia as a woody crop option for southern Australia, pp 204-214. Rural
Industries Research Development Corporation Publication No. 03/017, Canberra.

203. MASLIN B.R. & McDONALD M.W. (2004b) AcaciaSearch - Evaluation of Acacia as a woody crop
option for southern Australia. Rural Industries Research Development Corporation Publication No. 03/17,
Canberra.

204. MAUNDU P., KIBET S., MORIMOTO Y., IMBUMI M. & ADEKA R. (2009) Impact of Prosopis juliflora on
Kenya’s semi-arid and arid ecosystems and local livelihoods. Biodiversity 10(2-3):33-50.

205. Mc DERMOTT J.B. (1987) The control of Acacia saligna (Port Jackson Willow) by glyphosate in the South
Western Cape Province. Eighth National Weeds Congress, Potchefstroom.

206. Mc DONNELL M.J., STILES E.W., CHEPLICK G.P. & ARMESTO J.J. (1984) Bird dispersal of Phytolacca
americana L. and the influence of fruit removal on subsequent fruit development. American Journal
of Botany 71(7):895-901.

207. MELOCHE C. & MURPHY S.D. (2006) Managing Tree-of-Heaven (Ailanthus altissima) in Parks and
Protected Areas: A Case Study of Rondeau Provincial Park (Ontario, Canada). Environmental
Management 37(6):764-772.

208. MENDEL Z. (1998) Biogeography of Matsucoccus josephi (Homoptera: Matsucoccidae) as related

to host resistance in Pinus brutia and Pinus halepensis. Canadian Journal of Forest Research 28:323-330.
194
209. MIHULKA S., PYSEK P., MARTINKOVA J. & JAROSIK V. (2006) Invasiveness of Oenothera congeners alien
to Europe: Jack of all trades, master of invasion? Perspectives in Plant Ecology, Evolution and Systematics
8(2):83-96.

210. MILLAR M.A., BYRNE M., NUBERG I. & SEDGLEY M. (2008) A rapid PCR-based diagnostic test for the
identification of subspecies of Acacia saligna. Tree Genetics & Genomes 4(4):625-635.

211. MILLAR M. A., BYRNE M. & O’SULLIVAN W. (2011) Defining entities in the Acacia saligna (Fabaceae)
species complex using a population genetics approach. Australian Journal of Botany 59:137-148.

212. MILTON S.J. & HALL A.V. (1981). Reproductive biology of Australian Acacias in the south-western
cape province of South Africa. Transactions of the Royal Society of South Africa 44(3): 465-485.

213. MILTON S.J. & DEAN W.R.J. (1998) Alien plant assemblages near roads in arid and semi-arid South
Africa. Diversity & Distributions 4(4):175-187.

214. MILTON S.J., WILSON J.R.U., RICHARDSON D.M., SEYMOUR C.L., DEAN W.R.J., IPONGA D.M. &
PROCHEŞ Ş . (2007) Invasive alien plants infiltrate bird-mediated shrub nucleation processes in
arid savanna. Journal of Ecology 94(4):648-661.

215. MINISTRY OF AGRICULTURE & RURAL DEVELOPMENT - FLORA PROTECTION AND INSPECTION
SERVICES (2006) List of herbicides approved for sale in Israel. 82 pp. (In Hebrew).

216. MITCHELL D.S. & TUR N.M. (1975) The rate of growth of Salvinia molesta (S. auriculata Auct.) in
laboratory and natural conditions. Journal of Applied Ecology 12(1):213-225

217. MITCHELL A.A.,WILCOX D.G. & LAIDLAW (1988) Plants of the arid shrublands of Western Australia.
University if Western Australia Press, Medlands, Western Australia. 325 p.

218. MORAN V.C. & HOFFMANN J.H. (2011) Conservation of the fynbos biome in the Cape Floral Region:
the role of biological control in the management of invasive alien trees. BioControl in press.

219. MORRIS M.J. (1997) Impact of the gall-forming rust fungus Uromycladium tepperianum on the
invasive tree Acacia saligna in South Africa. Biological control 10:75-82.

220. MORRIS T.L., WITKOWSKI E.T.F. & COETZEE J.A. (2008) Initial response of riparian plant community
structure to clearing of invasive alien plants in Kruger National Park, South Africa. South African
Journal of Botany 74(3):485-494.
221. MOTARD E., MURATET A., CLAIR-MACZULAJTYS D., MACHON N. (2011) Does the invasive species
Ailanthus altissima threaten floristic diversity of temperate peri-urban forests? Comptes-Rendus
Biologies 334(12):872-879.

222. MOTOOKA P., CASTRO L., NELSON D., NAGAI G. & CHING L. (2003) Weeds of Hawai’i’s Pastures and
Natural Areas: An identification and Management Guide. College of Tropical Agriculture and Human
Resources, University of Hawai’i at Manoa.

223. MUYT A. (2001) Bush Invaders of South-East Australia: A guide to the identification and control of
environmental weeds found in South-East Australia. R.G & F.J. Richardson, pp.304.

224. NAHAL I. (1983) Le Pin Brutia (1ère partie). Forêt méditerranéenne 5(2):165-171.

225. NAHAL I. (1984) Le Pin Brutia (2ème partie). Forêt méditerranéenne 6(1):5-17.

226. NAKANO H., FUJIIY.,YAMADA K., KOSEMURA S.,YAMAMURA S., HASEGAWA K. & SUZUKIT. (2002) Isolation
and identification of plant growth inhibitors as candidate(s) for allelopathic substance(s), from
aqueous leachate from mesquite (Prosopis juliflora (Sw.) DC.) leaves. Plant Growth Regulation 37(2):
113-117.

227. NANDULA V.K., EUBANK T.W., POSTON D.H., CLIFFORD H., KOGER C.H. & REDDY K.N. (2006) Factors
affecting germination of horseweed (Conyza canadensis). Weed Science 54:898–902.

228. NASCIMBENE J. & MARINI L. (2010) Oak forest exploitation and black-locust invasion caused severe
shifts in epiphytic lichen communities in Northern Italy. Science of the Total Environment 408(22): 195
5506-5512.

229. NATIV R., EPHRATH J.R., BERLINER P. & SARANGA Y. (1999) Drought resistance and water use
efficiency in Acacia saligna. Australian Journal of Botany 47:577-586.

230. NICE G., JOHNSON B. & BAUMAN T. (2005) Pokeweed Control. Purdue University. Weed Science.
http://btny.purdue.edu/weedscience/2005/pokeweed05.pdf [accessed 17 February, 2010].

231. O’CONNOR T.G. (1995) Acacia karroo invasion of grassland: environmental and biotic effects
influencing seedling emergence and establishment. Oecologia 103(2):214-223.

232. O’SULLIVAN W., HUXTABLE D., GEORGE N. & BARTLE J. (2008) Review of Acacia saligna (Koojong
Wattle) and its potential for domestication for western australian dryland farming systems.
In Hobbs T.J., Barle J. & Bennell M. (Eds), Domestication potential of high priority species for woody biomass
crops in lower rainfall southern Australia, pp. 7-80, FloraSearch 3b. Rural Industries Research and Development
Corporation. p. 158.

233. OLD K.M. & DOS SANTOS CRISTOVAO C. (2003) A rust epidemic of the coffee shade tree
(Paraserianthes falcataria) in East Timor. In Agriculture: New Directions for a New Nation — East Timor
(Timor-Leste) edited by H. da Costa, C. Piggin, C.J. da Cruz and J. J. Fox, ACIAR Proceedings No. 113, 139-145.

234. OPPENHEIMER H.L. & BARTLETT R.T. (2002) New plant records from the main Hawaiian Islands.
Bishop Mus. Occas. Pap. 69(2):1-14.

235. ORR B.K. & RESH V.H. (1992) Influence of Myriophyllum aquaticum cover on Anophelus mosquito
abundance, oviposition, and larval microhabitat. Oecologia 90(4):474-482.

236. ORTAL R. & COHEN O. (1989) Rafts leaving Egypt. Land and Nature 31(4), (In Hebrew)

237. PACIFIC ISLAND ECOSYSTEMS at RISK (2008) Pennisetum clandestinum Hochst. Ex Chiov., Poaceae.
http://www.hear.org/pier/species/pennisetum_clandestinum.htm [accessed 20 July, 2009].

238. PALMER W.A., HEARD T.A., SHEPPARD A.W. (2010) A review of Australian classical biological control
of weeds programs and research activities over the past 12 years. Biological Control 52:271-287.

239. PARSONS W.T. & CUTHBERTSON E.G. (2001) Noxious weeds of Australia. Second edition. CSIRO
Publishing, Collingwood, Australia, p.698.

240. PASIECZNIK N.M., HARRIS P.J.C. & SMITH S.J. (2004) Identifying Tropical Prosopis Species: A Field
Guide. HDRA Publishing, Coventry, UK, p.30.
 241. PASIECZNIK N.M., OULD MOHAMED VALL A., NOURISSIER-MOUNTOU S., DANTHU P. (2006)
Discovery of a life history shift: precocious flowering in an introduced population of Prosopis.
Biological Invasions 8:1681-1687.

242. PETSIKOS C., DALIAS P. & TROUMBIS A.Y (2006) Effects of Oxalis pes-caprae L. invasion in olive
groves. Agriculture, Ecosystems & Environment 120:325-329.
243. PHELOUNG P.C., WILLIAMS P.A. & HALLOY S.R. (1999) A weed risk assessment model for use as a
biosecurity tool evaluating plant introductions. Journal of Environemental Management 57:239-251.
244. PIETERSE P. J. & McDERMOTT J. B. (1994) Season of application and glyphosate formulation as factors
influencing the efficacy of glyphosate on phyllode-bearing Australian acacias. South African Journal of
Plant and Soil 11(1):50-53.

245. PIETERSE A. H., KETTUNEN M., DIOUF S., NDAO I., SARR K., TARVAINEN A., KLOFF S. & HELLSTEN S.
(2003) Effective biological control of Salvinia molesta in the Senegal River by means of the weevil
Cyrtobagous salviniae. Ambio 32(7):458-462.

246. PLITMANN U., HEYN C.C., HELLER D. & SHMIDA A. (1983) Pictorial flora of Israel. Massada. 338 pp.
(In Hebrew)
247. POST J.A., KLEINJAN C.A., HOFFMANN J.H., IMPSON F.A.C. (2010) Biological control of Acacia
cyclops in South Africa: The fundamental and realized host range of Dasineura dielsi (Diptera:
Cecidomyiidae). Biological Control 53 68–75.
196
248. PRATT P.D. & T. D. CENTER T.D. (2011) Biocontrol without borders: the unintended spread of
introduced weed biological control agents. BioControl in press.
249. PRIEUR-RICHARD A.H., LAVOREL S., GRIGULIS K. & DOS SANTOS A. (2000) Plant community diversity
and invisibility by exotics: invasion of Mediterranean old fields by Conyza bonariensis and Conyza
canadensis. Ecology Letters 3:412-422.

250. PROCHE ., WILSON J.R.U., RICHARDSON D.M. & REJMANEK M. (2011) Native and naturalized range
size in Pinus: relative importance of biogeography, introduction effort and species traits. Global
Ecology & Biogeography in press.

251. PYŠEK P., RICHARDSON D.M., REJMANEK M., WEBSTER G.L., WILLIAMSON M. & KIRSCHNER J. (2004)
Alien plants in checklists and floras: towards better communication between taxonomists and
ecologists. Taxon 53(1):131-143.
252. PYŠEK P. & JAROSÍK V. (2005) Residence time determines the distribution of alien plants. In Inderjit
S., editor, Invasive plants: ecological and agricultural aspects, Basel: Birkhäuser Verlag, 77–96.
253. PYŠEK P., LAMBDON P.W., ARIANOUTSOU M., KÜHN I., PINO J. & WINTER M. (2008) Alien Vascular
Plants of Europe. In DAISIE, Handbook of Alien Species in Europe, p. 400, Springer Science, pp. 43-61.
254. PYŠEK P., RICHARDSON D.M. (2010) Invasive Species, Environmental Change and Management, and
Health. Annual Review of Environment and Resources 35:25-55.
255. PYŠEK P., BACHER S., CHYTRÝ M., JAROŠÍK V., WILD J., CELESTI-GRAPOW L., GASSÓ N., KENIS M.,
LAMBDON P.W., NENTWIG W., PERGL J., ROQUES A., SÁDLO J., SOLARZ W., VILÀ M. AND HULME P.E.
(2010) Contrasting patterns in the invasions of European terrestrial and freshwater habitats by
alien plants, insects and vertebrates. Global Ecology and Biogeography 19:317-331.
256. QUAYE M.,YAACOBY T. & RUBIN B. (2010) Biology and control of Heterotheca subaxillaris (camphor
weed) in Israel. Proceedings of the 2nd workshop on Invasive alien plants in Mediterranean-type regions of
the world, Trabzon, Turkey 02-06 August 2010, pp. 274-282.
257. QUEENSLAND GOVERNMENT (?) Beach primrose (Oenothera drummondii). Brochures & Information
sheets, Coasts, Plants, Coastal sand dunes: their vegetation and management , leaflet IV-05. p.2. http:
//www.epa.qld.gov.au/register/p00282aa.pdf [accessed 05 August, 2009]

258. QUEZEL P. & MEDAIL F. (2003) Ecologie et Biogéographie des forêts du bassin méditerranéen.
Elsevier. p. 571.
259. RAHAYU S., LEE S.S. & SHUKOR N.A.A (2010) Uromycladium tepperianum, the gall rust fungus from
Falcataria moluccana in Malaysia and Indonesia. Mycoscience 51:149–153.

260. RAMEAU J.C., MANSION D., DUME G. (1989) Flore forestière française - Tome 1: Plaines et collines.
Institut pour le Développement Forestier, p.1785.

261. RANDALL J.M. & MARINELLI J. (1996) Invasive plants: weeds of the global garden. Brooklyn Botanic
Garden Publications p. 111.

262. RANDALL J. (1999) Import Risk Analysis: Importation of weed species by live animals and
unprocessed fibre of sheep and goats. Regulatory Authority, Ministry of Agriculture and Forestry.
Wellington, New-Zealand. p. 25.

263. RAVIV V. (1991) Plants and Animals of the Land of Israel. Azaria Alon, Ed. Volume 9 Flowerless Plants.
Third Edition. 259 pp. (In Hebrew)

264. REICHARD S. (2001) The search for patterns that enable prediction of invasion. In Groves R.H.,
Panetta F.D. & Virtue J.G. (Eds) Weed Risk Assessment, CSIRO publishing, pp. 10-19.

265. REJMÁNEK, M., RICHARDSON, D.M., HIGGINS, S.I., PITCAIRN, M.J. AND GROTKOPP, E. (2005) Ecology of
invasive plants: state of the art. In Mooney, H.A., Mack, R.M., McNeely, J.A., Neville, L., Schei, P. and Waage,
J., editors, Invasive alien species: searching for solutions, Washington, DC: Island Press, 104–161.

266. REMALEY T. & SWEARINGEN J.M. (2005) Fact sheet: White Poplar. Plant Conservation Alliance’s Alien
Plant Working Group. p.3. 197
267. REVES Y. (2008) A Guide for Eucalyptus Species in Israel. Second edition. Gestelit publishing house, Haifa,
Israel, 245 pp. (in Hebrew and in English)

268. RICE S.K., WESTERMAN B. & FEDERICI R. (2004) Impacts of the exotic, nitrogen-fixing black locust
(Robinia pseudoacacia) on nitrogen cycling in a pine-oak ecosystem. Plant Ecology 174:97–107

269. RICHARDSON D.M. (1998) Forestry trees as invasive aliens. Conservation Biology 12(1):18-26.

270. RICHARDSON D.M. (2001) Plant Invasions. In: Levin, S. (ed.), Encyclopedia of Biodiversity, Volume 4.,
pp. 677-688. Academic Press, San Diego.

271. RICHARDSON D.M., PYŠEK P., REJMÁNEK M., BARBOUR M.G., PANETTA F.D. & WEST C.J. (2000)
Naturalization and invasion of alien plants: concepts and definitions. Diversity and Distributions
6:93-107.

272. RICHARDSON D.M., ALLSOPP N., D’ANTONIO C.M., MILTON S. & REJMANEK M. (2000) Plant invasions
– the role of mutualisms. Biological Review 75:65-93.

273. RICHARDSON D.M. VAN WILGEN B.W. (2004) Invasive alien plants in South Africa: how well do we
understand the ecological impacts? South African Journal of Science 100:45-52.

274. RICHARDSON D.M., ROUGET M., RALSTON S.J., COWLING R.M., VAN RENSBURG B.J. & THUILLIER
W. (2005) Species richness of alien plants in South Africa: Environmental correlates and the
relationship with indigenous plant species richness. Ecoscience 12(3):391-402.

275. RICHARDSON D.M. & PYŠEK P. (2006) Plant invasions: Merging the concepts of species invasiveness
and community invasibility. Progress in Physical Geography 30(3):409-431.

276. RICHARDSON D.M., HOLMES P.M., ESLER K.J., GALATOWITSH S.M., STROMBERG J.C., KIRKMAN
S.P., PYSEK P. & HOBBS R.J. (2007) Riparian vegetation: degradation, alien plant invasions, and
restoration prospects. Diversity and Distributions 13:126-139.

277. RICHARDSON D.M. & THUILLER W. (2007) Home away from home - objective mapping of high-risk
source areas for plant introductions. Diversity & Distributions 13(3):299-312.

278. RICHARDSON D.M. & KLUGE R.L. (2008) Seed banks of invasive Australian Acacia species in South
Africa: Role in invasiveness and options for management. Perspectives in Plant Evolution, Ecology and
Systematics 10(3):161-177.
 279. RICHARDSON D.M. & REJMÁNEK M. (2011) Trees and shrubs as invasive alien species – a global
review. Diversity & Distributions 17:788-809.

280. RICHARDSON D.M., PYŠEK P. & CARLTON J.T. (2011) A compendium of essential concepts and
terminology in invasion ecology. In Richardson D.M. (Ed) Fifty Years of Invasion Ecology:The Legacy of Charles
Elton. First edition, Blackwell Publishing Ltd. pp.409-420.

281. RICHARDSON F.J. & RICHARDSON R.G. & SHEPHERD R.C.H (2011) Weeds of the South-East – An
identification guide for Australia. Second edition. R.G & F.J. Richardson Meredith,Victoria, Australia.

282. RICHARDSON R.G. & MCKENZIE D.N. (1981) Regeneration of, and toxicity of 2,4-D, to root
fragments of silver-leaf nightshade (Solanum elaeagnifolium Cav.). Journal of the Australian Institute of
Agriculture Science 47:48-50.

283. RIZVI S.J.H. & RIZVI V. (1992) Allelopathy: Basic and applied aspects. Chapman & Hall, London, p. 504.

284. ROILOA S.R., RODRÍGUEZ-ECHEVERRÍA S., DE LA PEÑA E. & FREITAS H. (2010) Physiological
integration increases the survival and growth of the clonal invader Carpobrotus edulis. Biological
Invasions 12:1815-1823.

285. ROIRON P.,ALI A.A., GUENDON J.L., CARCAILLET C.& TERRAL J.F. (2004) Preuve de l’indigénat de
Populus alba L. dans le Bassin méditerranéen occidental. Comptes Rendus Biologies 327(2):125-132.

286. ROMANOWSKI N. (2011) Wetland weeds, causes, cures and compromises. CSIRO publishing,
198 Australia, p. 140.

287. RUTHERFORD M.C. & POWRIE L. (1993) Allelochemic control of biomass allocation in interacting
shrub species. Journal of Chemical Ecology 19(5):1573-1561.

288. SADE R.,YAACOBY T. & RUBIN B. (2007) Crownbeard (Verbesina encelioides): physiology, geographical
distribution and response to herbicides in Israel. Novel and Sustainable Weed Management in Arid and
Semi-Arid Ecosystems, Rehovot, Israel, October 2007. Conferences Abstract, p.41.

289. SAWER E.R. (1922) Report on forestry and agriculture in Palestine. A Review of the agricultural
situation in Palestine. Department of Agriculture and Fisheries. Palestine.

290. SAWER E.R. (1931) Report of the department of agriculture and forests for the years 1927-1930.
p. 167.

291. SCHUELLER S.K. (2004) Self-pollination in island and mainland populations of the introduced
Hummingbird-pollinated plant, Nicotiana glauca (Solanaceae). American Journal of Botany 91(5):
672-681.

292. SFORZA R. & JONES W.A. (2007) Potential for classical biocontrol of silverleaf nightshade in the
Mediterranean Basin. Bulletin OEPP/EPPO Bulletin 37:156-162.

293. SHALMON B. (1993) A field guide to the land mammals of Israel their tracks and signs. Keter.
216 pp. (In Hebrew)

294. SHARMA G.P., RAGHUBANSHI A.S. & SINGH J.S. (2005) Lantana invasion: An overview. Weed Biology and
Management 5(4):157-165.

295. SHARMA G.P. & RAGHUBANSHI A.S. (2009) Lantana invasion alters soil nitrogen pools and processes
in the tropical dry deciduous forest of India. Applied Soil Ecology 42(2):134-140.

296. SHETTY K.G., MINNIS A.M., ROSSMAN A.Y., JAYACHANDRAN K. (2011) The Brazilian peppertree
seed-borne pathogen, Neofusicoccum batangarum, a potential biocontrol agent. Biological Control
56:91-97.

297. SHIELDS E.J., DAUER J.T., VANGESSEL M.J. & NEUMANN G. (2006) Horseweed (Conyza canadensis)
seed collected in the planetary boundary layer. Weed Science 54(6):1063-1067.

298. SHLUKER A. (2002) Verbesina encelioides [(Cav.) Bentham & Hooker fil. ex Gray] ssp. exauriculata
[Robinson & Greenman]. Harmful Non-Indigenous Species (HNIS) Report, Hawaiian Ecosystems at Risk
project (HEAR). p.12
299. SHMIDA A. (1986) Handbook of wildflowers of Israel: Mediterranean Flora. Keter. 308 pp.
(In Hebrew).

300. SHMIDA A. (1992) Handbook of Trees and Bushes of Israel. Keter. 328 pp. (In Hebrew)

301. SHMIDA A. (1992) Handbook of wildflowers of Israel: Desert Flora. Keter. 334 pp. (In Hebrew).

302. SHMIDA A. (2005) MAPA’s dictionary of plants and flowers in Israel. Mapa. 493 pp. (In Hebrew)

303. SHMIDA A. & POLAK G. (2007) Red data book: Endangered plants of Israel, Vol. 1. Published by the
Nture & Parks Authority, 492 pp. (In Hebrew).

304. SHRESTHA A., HEMBREE K. & WRIGHT S. (2008) Biology and Management of horseweed and hairy
fleabane in California. University of California, Division of Agriculture and Natural Resources. Publication
8314. p.9.

305. SILVA S.I., BOZINOVIC F. & JAKSIC F.M. (2005) Frugivory and seed dispersal by foxes in relation to
mammalian prey abundance in a semiarid thornscrub. Austral Ecology 30:739–746.

306. SILVA S.I., JAKSIC F.M. & BOZINOVIC F. (2005) Nutritional ecology and digestive response to dietary
shift in the large South American fox, Pseudalopex culpaeus. Revista Chilena de Historia Natural
78:239-246.

307. SIMBERLOFF D. (1996) How risky is biological control ? Ecology 77(7): 1965-1974.

308. SIMBERLOFF D. (2012) Risks of biological control for conservation purposes. BioControl 57(2):
199
263-276.

309. SINAI I., CHAZAN G. & KOREN S. (2011) Control of invasive plants in the Na’aman and the Tel Afek
Nature Reserves. Proceedings of the Nature & Parks Authority annual conference, Herzelya 28 December
2011. p.16. (In Hebrew).

310. SINGH M., TAMMA R.V. & NIGG H.N. (1989) HPLC identification of allelopathic compounds from
Lantana camara. Journal of Chemical Ecology 15(1):81-89.

311. SOMERVILLE S., SOMERVILLE W. & COYLE R. (2011) Regenerating native forest using splatter gun
techniques to remove Lantana. Ecological Management & Restoration 12(3), 164-174.

312. SPONNER A. (2007) Oenothera drummondii Hook. Florabase The Western Australian Flora. Western
Australia Government. http://florabase.calm.wa.gov.au/browse/profile/6138 [accessed 05 August, 2009].

313. STARR F., STARR K. & LOOPE L. (2003) Ficus benghalensis Indian banyan tree - Moraceae. US Geological
Survey, Biological Resources Division, Haleakala Field Station, Maui, Hawai’i, p. 5.

314. STARR F., STARR K. & LOOPE L. (2003) Ficus microcarpa Chinese banyan - Moraceae. US Geological
Survey, Biological Resources Division, Haleakala Field Station, Maui, Hawai’i, p. 8.

315. STARR F., STARR K. & LOOPE L. (2003) Ficus religiosa Bo Tree - Moraceae. US Geological Survey,
Biological Resources Division, Haleakala Field Station, Maui, Hawai’i, p. 4.

316. STATE of WASHINGTON, DEPARTMENT OF ECOLOGY (2003) Parrotfeather (Myriophyllum

aquaticum) - Technical information. www.ecy.wa.gov/programs/wq/plants/weeds/parrot.htm/invasive.org/
weedus/ubject.html?sub=3054 [accessed 16 August, 2009].

317. STOREY R. & JONES R.G.W (1979) Response of Atriplex spongiosa and Suaeda monoica to salinity.
Plant Physiology 63:156-162.

318. STRYDOM M., ESLER K.J. & WOOD A.R. (2011) Acacia saligna seed banks: Sampling methods and
dynamics, Western Cape, South Africa. South African Journal of Botany, in press.

319. SULLIVAN P.R., POSTLE L.A., JULIEN M. (2011) Biological control of Salvinia molesta by Cyrtobagous
salviniae in temperate Australia. Biological Control 57:222–228.

320. TAKESHI T., SHIGENOBU T., NORIKAZU Y. & KAZUYOSHI F. (2007) Inhibition of the regeneration of
Japanese black pine (Pinus thunbergii ) by black locust (Robinia pseudoacacia) in coastal sand
dunes. Journal of Forest Research 12(5):350-357.
 321. TALEB A., BOUHACHE M. & EL MFADI B. (2010) Nouvelle espèce menaçant la biodiversité au Maroc:
Verbesina encelioides (Asteraceae). Proceedings of the 2nd international workshop on invasive plants in
the Mediterranean type regions of the world,Trabzon,Turkey, 02-06 August 2010, European and Mediterranean
Plant Protection Organization (EPPO).

322. TANJI A., BOULET C. & HAMMOUMI M. (1984) Contribution to the study of the biology of Solanum
elaeagnifolium Cav. (Solanaceae), a weed of crops in the irrigated perimeter of the Tadla
(Morocco). Weed Research 24:401-409.

323. TASSIN J., RIVIERE J.N. & CLERGEAU P. (2007) Reproductive versus Vegetative Recruitment of the
Invasive Tree Schinus terebenthifolius: Implications for Restoration on Reunion Island. Restoration
Ecology 15(3):412-419.

324. THOMPSON C.R. & HABECK D.H. (1989) Host specificity and biology of the weevil Neohydronomus
affinis [Coleoptera: Curculionidae] a biological control agent of Pistia stratiotes. BioControl
34(3):299-306.

325. THOMPSON G.D., ROBERTSON M.P., WEBBER B.L., RICHARDSON D.M., LE ROUX J.J. & WILSON J.R.U.
(2011) Predicting the subspecific identity of invasive species using distribution models: Acacia
saligna as an example. Diversity & Distributions 17:1001-1014.

325a THOMPSON G.D., BELLSTEDT D.U., BYRNE M., MILLAR M.A., RICHARDSON D.M., WILSON J.R.U. & LE
ROUX J.J. (2012) Cultivation shapes genetic novelty in a globally important invader. Molecular Ecology
200 21:3187-3199.

325. THOMPSON K. & DAVIS M.A. (2011) Why research on traits of invasive plants tells us very little.
Trends in Ecology & Evolution 26(4):155-156.

326. TOTLAND Ø., NYEKO P., BJERKNES A.L., HEGLAND S.J. & NIELSEN A. (2005) Does forest gap size affect
population size, plant size, reproductive success and pollinator visitation in Lantana camara, a
tropical invasive shrub ? Forest Ecology and Management 215:329-338.

327. TOZER M.G. (1998) Distribution of the soil seedbank and influence of fire on seedling emergence
in Acacia saligna growing on the central coast of New South Wales. Australian Journal of Botany
46:743-755.

328. TROPICAL FORAGES (?) Pennisetum clandestinum. http://www.tropicalforages.info/key/Forages/Media/

Html/Pennisetum_clandestinum.htm

329. TUCKER G.C. (1994) Revision of the Mexican Species of Cyperus L. (Cyperaceae). Systematic Botany
Monographs, vol. 43, 213p.

330. TYE D.R.C. & DRAKE D.C. (2011) An exotic Australian Acacia fixes more N than a coexisting
indigenous Acacia in a South African riparian zone. Plant Ecology, in press.

331. UNITED STATES DEPARTMENT OF AGRICULTURE (2009) Agricultural Research Service (ARS), National
Genetic Resources Program. Germplasm Resources Information Network – (GRIN). Online database. http:
//www.ars-grin.gov/cgi-bin/npgs/html/taxon.pl?400375 [accessed 02 August, 2009].

332. VAN DRIESCHE R.G., CARRUTHERS R.I., CENTER T., HODDLE M.S., HOUGH-GOLDSTEIN J., MORIN
L., SMITH L., WAGNER D.L. et al. (2010) Classical biological control for the protection of natural
ecosystems. Biological Control 54 Supplement 1:S2-S33.

333. VAN KLEUNEN M., FISHER M. & JOHNSON S.D. (2007) Reproductive assurance through self-
fertilization does not vary with population size in the alien invasive plant Datura stramonium.
Oikos 116(8):1400-1412.

334. VAN WILGEN B.W., RICHARDSON D.M., LE MAITRE D.C., MARAIS C. & MAGADLELA D. (2001)
The economic consequences of alien plant invasions: Examples of impacts and approaches to
sustainable management in South Africa. Environment, Development and Sustainability 3:145-168.

335. VARNHAM K. (2006) Non-native species in UK overseas territories – A review. Joint Nature
Conservation Committee No 372, Peterborough, p.39.
336. VELDTMAN R., LADO T.F., BOTES A., PROCHES S.¸ TIMM A.E., GEERTSEMA H. & CHOWN S.L. (2011)
Creating novel food webs on introduced Australian acacias: indirect effects of galling biological
control agents. Diversity & Distributions 17:958-967.

337. VILÀ M., BARTOMEUS I., GIMENO I., TRAVESET A. & MORAGUES E. (2006) Demography of the Invasive
Geophyte Oxalis pes-caprae Across a Mediterranean Island. Annals of Botany 97:1055-1062.

338. WEAVER S.E. (2001) The biology of Canadian weeds. 115. Conyza canadensis. Canadian Journal of Plant
Science 81(4):867-875.

339. WEBER E. (2003) Invasive plant species of the world, a reference guide to environmental weeds.
CABI Publishing, Oxon, p.548.

340. WERSAL R.M. & MADSEN J.D. (2011) Comparative effects of water level variations on growth
characteristics of Myriophyllum aquaticum. Weed Research 51(4):386-393.

341. WEST J.G. (1984) A revision of Dodonaea Miller (Sapindaceae) in Australia. Brunonia 7:1–194.

342. WHITNEY K.D & GABLER C.A. (2000) Rapid evolution in introduced species, ‘invasive traits’ and
recipient communities: challenges for predicting invasive potential. Diversity and Distributions
14:569-580.

343. WILEN C.A. & HOLT J.S. (1996) Physiological mechanisms for the rapid growth of Pennisetum
clandestinum in Mediterranean climates. Weed Research 36(3):213-225.
201
344. WILLIAMSON M. (1996) Biological invasions. Population and community biology series 15, Chapman &
Hall, London. p.244.

345. WILLIAMSON M. & FITTER A. (1996) The varying success of invaders. Ecology 77(6):1661-1666.

346. WILLIS A.J. & MEMMOTT J. (2005) The potential for indirect effects between a weed, one of its
biocontrol agents and native herbivores: A food web approach. Biological Control 35:299-306.

347. WILSON E.O (2003) The future of life. Abacus, London, p. 230.

348. WILSON J.R.U., REES M. & AJUONU O. (2006) Population regulation of a classical biological control
agent: larval density dependence in Neochetina eichhorniae (Coleoptera: Curculionidae), a
biological control agent of water hyacinth Eichhornia crassipes. Bulletin of Entomological Research 96:
145–152.

349. WILSON J.R.U., AJUONU O., CENTER T.D., HILL M.P., JULIEN M.H., KATAGIRA F.F., NEUENSCHWANDER
P., NJOKA S.W., OGWANG J., REEDER R.H. & VAN T. (2007) The decline of water hyacinth on Lake
Victoria was due to biological control by Neochetina spp. Aquatic Botany 87:90-93.

350. WILSON J.R.U, GAIRIFO C., GIBSON M.R., ARIANOUTSOU M., BAKAR B.B., BARET S., CELESTI-GRAPOW
L., DI TOMASO J.M., DUFOUR-DROR J.M., KUEFFER C., KULL C.A., HOFFMANN J.H., IMPSON F.A.C.,
LOOPE L.L., MARCHANTE E., MARCHANTE H., MOORE J.L., MURPHY D.J., TASSIN J., WITT A., ZENNI
R.D., RICHARDSON D.M. (2011) Risk assessment, eradication, containment, and biological control:
global efforts to manage Australian acacias before they become widespread invaders. Diversity &
Distribution 17:1030-1046.

351. WITTENBERG R. & COCK M.J.W. (2001) Invasive alien species: A toolkit of best prevention and
management practices. Global Invasive Species Programme, CABI publishing, Oxon, p.240.

352. WOOD A.R. & MORRIS M.J. (2007) Impact of the gall-forming rust fungus Uromycladium
tepperianum on the invasive tree Acacia saligna in South Africa: 15 years of monitoring. Biological
Control 41:68-77.

353. YAACOBY T. (2007) Invasive alien species (IAS) current situation in Israel. Novel and Sustainable
Weed Management in Arid and Semi-Arid -Ecosystems, Rehovot, Israel, October 2007. Conference abstracts,
p.42.

354. YAACOBY T. & SEPLYARSKY V. (2011) Epiblema strenuana (Walker, 1863) (Lepidoptera:Tortricidae),
a new species in Israel. EPPO Bulletin 41(2):243-246.
 355. YEATES A.G. & SCHOOLER S.S. (2011) Influence of Lantana camara and its removal on tree
dynamics in a recently burnt wet Sclerophyll forest in Northern NSW. Ecological Management &
Restoration 12(3), 236-241.

356. YELENIK S.G., STOCK W.D. & RICHARDSON D.M. (2004) Ecosystem level impacts of invasive Acacia
saligna in the South African fynbos. Restoration Ecology 12(1):44-51.

357. ZAFRIR D. (1976) Ornamental flowers in Israel, trees, shrubs, dwarf shrubs and climbers. Am Oved
– Culture and Education, 143 pp. (In Hebrew).

358. ZALUCKI M.P., DAT M.D. & PLAYFORD J. (2007) Will biological control of Lantana camara ever
succeed? Patterns, processes & prospects. Biological Control 42(3):251-261.

359. ZAPLATA M.K.,WINTER S., BIEMELT D. & FISCHER A. (2011) Immediate shift towards source dynamics:
The pioneer species Conyza canadensis in an initial ecosystem. Flora 206(11):921-927.

360. ZENNI R.D., WILSON J.R.U, LE ROUX J.J. & RICHARDSON D.M. (2009) Evaluating the invasiveness of
Acacia paradoxa in South Africa. South African Journal of Botany, in press.

361. ZIMMELS Y., KIRZHNER F. & MALKOVSKAJA A. (2006) Application of Eichhornia crassipes and Pistia
stratiotes for treatment of urban sewage in Israel. Journal of Environmental Management 81(4):
420-428.

362. ZIMMERMANN H.G., HOFFMANN J.H. & WITT A.B.R. (2006) A South African Perspective on Prosopis.
202 Biocontrol News & Information 27(1):6-10.
 Appendix
List of Alien Plant Species in Natural and
Disturbed Areas in Israel
Prepared with Dr. Ori Fragman-Sapir, head scientist of the
Jerusalem Botanical Gardens, the Hebrew University of Jerusalem.

Selection of Species Included in the Table

The aim of this table is to propose a first comprehensive database of alien plant species
found in Israel in natural and disturbed habitats. All the plant species in the table are
alien species according to the definitions of Pyšek et al. (2004). Taxa that were recorded
at least three times, with at least one of the observations having occurred during the 203
past 20 years (since 1990), were included in the table. Alien plant species in aquatic
habitats were included in the table, even if they were observed only once, due to their
high invasive potential.

Alien species grown in Israel that were not observed outside their area of cultivation, as
well as alien species that do not reproduce, either sexually or asexually, in the conditions
found in Israel were not included in the table. Similarly, alien species for which there
is no information, whether from the National Herbarium at the Hebrew University of
Jerusalem or any other sources were not listed in the database.

Presence in Israel and Origin

Three basic regions in Israel were distinguished for determining species presence:
(a) the Mediterranean, (b) semi-arid and (c) desert regions.

The alien species areas of origin were divided as follows:

Continent Region Sub-region Code in table

America Am
North America AmN
Temperate North America AmNTemp
Arid North America AmNArid
Central America AmC
South America AmS
Tropical South America AmSTrop
Temperate South America AmSTemp
 Continent Region Sub-region Code in table
Africa Af
Mediterranean Africa AfMed
Tropical Africa AfTrop
Eastern Tropical Africa AfTropE
Southern Tropical Africa AfTropS
Cape Region AfCape
Europe Eu
Central and Northern Europe EuCont
Mediterranean Europe EuMed
Western Mediterranean Europe EuMedW
Eastern Mediterranean Europe EuMedE
Australia Au
Northern Australia AuN
204 Arid Australia AuArid
Western Arid Australia AuAridW
Central Arid Australia AuAridC
Eastern Arid Australia AuAridE
Southern Australia AuS
Southwestern Australia AuSW
Southeastern Australia AuSE
Asia As
Temperate Asia AsTemp
Western Temperate Asia AsTempW
Central Temperate Asia AsTempC
Arabian Peninsula AsArP
Tropical Asia AsTrop
India AsIndia
Southeastern Asia AsSE

Incidence and Habitat

Four degrees of incidence were determined, the first two quantitative, based on the
frequency scale of Fragman et al. (1999) and the other two qualitative:

Very rare (species observed in less than 20 sites in Israel, with each site up to
1 sq.km.)

Rare (species observed in 20-1000 sites in Israel, each site up to 1 sq. km)

Common

Very Common
It should be noted that incidence is evaluated on a country-wide scale and expresses
the spatial distribution in Israel. Thus, a plant species, all of whose individuals are
spread only over only a few square kilometers, will be considered “very rare”, even if
on a local scale it forms dense units with many individuals, that can cause significant
ecological damage. At the same time, a plant species with many representatives
distributed over extensive areas in the country will be considered “common” even if
its representatives do not form dense concentrations.

The following habitats populated by alien species in Israel were determined:

Habitat Type Code in table ¿

Natural dry habitat (undisturbed) NDL

Natural moist habitat (undisturbed) NWL

Disturbed Dry Habitat DDL

205
Cultivated fields, parks, gardens DDL agri

Planted forests DDL plfor

Roadsides, railroad tracks, infrastructure DDL rdsid

Garbage dumps, landfills DDL refsi

Disturbed or artificial moist habitats DWL

Urban environment UE
 206
List of Alien Plant Species in Natural and Disturbed Areas in Israel

Scientific Name Origin Presence in Israel Habitat Occurrence

1 Abutilon theophrastii AsSE, AsTempC Mediterranean DDHagri rare

2 Acacia cyclops AuSW Mediterranean NDH rare
3 Acacia farnesiana AmC Mediterranean NDH,DDHagri,DDHrdsid common
4 Acacia karroo AfTropS, AfTropE Mediterranean DDHrdsid very rare
5 Acacia paradoxa AuSE Mediterranean DDHagri,DDHrdsid very rare
6 Acacia salicina AuAridE, AuSE desert NDH,DDHplfor,DDHrdsid,UE common
7 Acacia saligna AuSW Mediterranean, semi-arid NDH,NWH,DDHall,DWH very common
8 Acacia sclerosperma AuAridW semi-arid DDHrdsid very rare
9 Acacia victoriae AuArid Mediterranean, semi-arid DDHplfor,DDHrdsid rare
NDH, DDHagri, DDHplfor,
10 Achyranthes aspera AsSE Mediterranean, desert common
DDHrdsid
11 Agave americana AmNArid Mediterranean, semi-arid, desert DDHagri, DDHrdsid common
NDH,DDHagri,DDHplfor,DDHrd
12 Ailanthus altissima AsSE Mediterranean common
sid,UE
13 Amaranthus albus AmNTemp Mediterranean, semi-arid, desert DDHagri,DDHrdsid,DDHrefsit,UE common
14 Amaranthus blitoides AmN Mediterranean, semi-arid, desert DDHall DWH, UE very common
15 Amaranthus blitum EuCont, AsTemp Mediterranean, semi-arid, desert DDHall UE very common
 Scientific Name Origin Presence in Israel Habitat Occurrence

16 Amaranthus cruentus AmC, AmSTrop Mediterranean, semi-arid, desert DDHagri, DDHrdsid, DWH common
17 Amaranthus graecizans ? Mediterranean, semi-arid, desert DDHagri, DDHrdsid, UE very common
18 Amaranthus muricatus AmS Mediterranean DDHagri,DDHrefsit very rare
19 Amaranthus palmeri AmNArid Mediterranean, semi-arid, desert DDHagri, DWH common
20 Amaranthus retroflexus AmNTemp Mediterranean, semi-arid, desert DDHagri,DDHrdsid,DDHrefsit common
21 Amaranthus rudis AmNTemp Mediterranean NWH,DWH rare
22 Amaranthus spinosus AmC, AmSTrop Mediterranean, semi-arid DDHagri,DDHrdsid rare
23 Amaranthus viridis AmC, AmSTrop Mediterranean DDHagri,DDHrdsid,DDHrefsit very common
24 Ambrosia confertiflora AmNArid Mediterranean NWH,DDHrdsid very rare
25 Ambrosia tenuifolia AmC, AmSTrop Mediterranean DDHrdsid,DDHrefsit very rare
26 Anacyclus radiatus EuMedW, AfMed Mediterranean DDHrdsid very rare
27 Anoda cristata AmNArid, AmC, AmS Mediterranean, semi-arid DDHagri rare
28 Antirrhinum siculum EuMedW Mediterranean UE very rare
29 Araujia sericifera AmSTrop Mediterranean DDHagri rare
30 Argremone mexicana AmNArid Mediterranean, semi-arid, desert DDHagri,DDHrdsid very rare
31 Artemisia arborescens EuMedW Mediterranean NDH, DDHagri, DDHrdsid very rare
32 Artemisia scoparia Eu, AsTemp Mediterranean, desert DDHagri very rare
33 Asparagus setaceus AfTropS Mediterranean DDHagri common

NWH, DDHagri DDHplfor,

34 Aster subulatus AmNTemp, AmC Mediterranean, semi-arid, desert very common
DDHrdsid, DWH
207
 208
Scientific Name Origin Presence in Israel Habitat Occurrence

35 Atriplex holocarpa AuArid desert NDH,DDHrdsid common

36 Atriplex muelleri AuSE, AuAridC desert DDHrdsid, UE very rare

37 Atriplex semibaccata AuArid Mediterranean, semi-arid, desert DDHagri, DDHrdsid, DWH common

38 Atriplex suberecta AuSW, AuAridW Mediterranean, semi-arid, desert DDHagri, DDHrdsid, DWH rare

39 Azolla filiculoides Am Mediterranean NWH, DWH very rare

40 Bassia scoparia EuCont, AsTemp Mediterranean, semi-arid DDHrdsid rare

DDHagri, DDHplfor,
41 Bidens pilosa AmSTrop Mediterranean, semi-arid, desert common
DDHrdsid,DDHrefsit

42 Borrichia frutescens AmC Mediterranean UE very rare

43 Brassica napus EuMedW Mediterranean DDHagri rare

44 Bromus catharticus AmS Mediterranean, semi-arid, desert DDHagri, DDHrdsid common

45 Carpobrotus acinaciformis AfCape Mediterranean NDH,UE common

46 Carpobrotus edulis AfCape Mediterranean NDH,UE common

47 Cenchrus echinatus AmC, AmSTrop Mediterranean, semi-arid DDHagri, DDHrdsid common

48 Cenchrus incertus AmN, AmC, AmSTrop Mediterranean DDHagri rare

49 Centranthus ruber susp. ruber EuMed, AfMed Mediterranean DDHagri, DDHrdsid, UE rare

50 Chenopodium ambrosioides AmC, AmSTrop Mediterranean, semi-arid, desert DDHagri, DDHrdsid, UE common

51 Chenopodium missouriense AmNTemp Mediterranean DDHrdsid, UE rare

52 Chenopodium urbicum EuCont, AsTemp Mediterranean DWH rare

 Scientific Name Origin Presence in Israel Habitat Occurrence

53 Chloris gayana AfTrop Mediterranean, semi-arid, desert DDHagri, DDHrdsid common

54 Chloris pycnothrix AfTrop Mediterranean, semi-arid DDHagri very rare

55 Chloris virgata AmSTrop, AsTrop Mediterranean, semi-arid, desert DDHagri common

56 Ciclospermum leptophyllum AmC, AmSTrop Mediterranean NDH, DDHagri, DDHrdsid very rare

57 Cirsium vulgare EuCont, AsTemp Mediterranean DDHrdsid, DWH very rare

NWH, DDHagri, DDHplfor,

58 Conyza albida AmSTrop Mediterranean, semi-arid, desert rare
DDHrdsid, DWH

NWH, DDHagri, DDHplfor,

59 Conyza bonariensis AmSTrop Mediterranean, semi-arid, desert very common
DDHrdsid, DWH, UE

NWH, DDHagri, DDHplfor,

60 Conyza canadensis AmN Mediterranean, semi-arid, desert very common
DDHrdsid, DWH, UE

61 Coronopus didymus AmSTrop Mediterranean UE common

62 Cuscuta campestris AmN Mediterranean, semi-arid, desert DDHagri rare

AmC, AmSTrop,
63 Cyperus imbricatus Mediterranean DWH very rare
AsTrop, AfTrop

64 Cyperus involucratus AfTropE Mediterranean, semi-arid NWH very rare

AmN, AmC, AmSTrop,

65 Cyperus odoratus Mediterranean NWH rare
AsTrop, AfTrop

66 Datura ferox AsTrop Mediterranean DDHagri, DDHrdsid rare

67 Datura innoxia AmC, AmSTrop Mediterranean, semi-arid, desert DDHall DWH, UE very common

68 Datura stramonium AmC, AmSTrop Mediterranean, semi-arid, desert DDHall DWH, UE very common
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Scientific Name Origin Presence in Israel Habitat Occurrence

69 Diospyros virginiana AmN Mediterranean NWH, DWH very rare

70 Diplotaxis tenuifolia EuCont, EuMed, AfMed Mediterranean UE very rare

AmC, AmSTrop,
71 Dodonaea viscosa Mediterranean NDH, DDHrdsid rare
AsTrop, AfTrop, Au

72 Eclipta prostrata AmN, AsTemp Mediterranean NWH, DDHagri, DWH common

73 Ehrharta erecta AfTropS Mediterranean DDHagri very rare

74 Eichhornia crassipes AmSTrop Mediterranean NWH, DWH very rare

75 Einadia nutans AuS, AuSW desert DDHagri very rare

76 Eleusine indica AfTrop Mediterranean, semi-arid, desert DDHagri, DDHrdsid common

77 Elodea canadensis AmN Mediterranean NWH, DWH rare

78 Enchylaena tomentosa Au Mediterranean, semi-arid, desert DDHrdsid, UE rare

79 Eragrostis echinochloidea AfTropS desert DDHagri very rare

80 Eragrostis japonica Af, AmN, AsTemp Mediterranean NWH, DWH very rare

81 Eragrostis palmeri AmNArid, AmC Mediterranean DDHagri, DDHrdsid rare

82 Eragrostis sarmentosa AfTropS, AfCape Mediterranean, desert NWH, DWH very rare

83 Eucalyptus camaldulensis Au Mediterranean, semi-arid NDH, NWH, DWH rare

84 Euphorbia gramineae AmC Mediterranean DDHagri,DDHrdsid very rare

85 Euphorbia heterophylla AmNArid Mediterranean, semi-arid DDHagri, DDHrdsid common

 Scientific Name Origin Presence in Israel Habitat Occurrence

86 Euphorbia hirta AmC Mediterranean, semi-arid, desert DDHagri, DDHrdsid common

DDHagri, DDHplfor, DDHrdsid,

87 Euphorbia lasiocarpa AmN Mediterranean, semi-arid, desert very common
DWH

88 Euphorbia maculata AmN Mediterranean, desert DDHagri rare

89 Euphorbia prostrata AmC, AmSTrop Mediterranean, semi-arid, desert DDHagri, UE common

90 Euphorbia serpens AmC, AmSTrop Mediterranean, semi-arid, desert DDHagri, UE common

91 Euphorbia supina AmN Mediterranean, semi-arid, desert DDHagri, UE rare

Eu, AsTemp, AsTrop,

92 Fallopia convolvulus Mediterranean, semi-arid DDHagri, UE very rare
AfMed

93 Ficus benghalensis AsIndia semi-arid NWH very rare

94 Ficus religiosa AsIndia, AsTrop semi-arid NWH very rare

95 Ficus retusa AsTrop, AuN semi-arid NWH very rare

96 Galinsoga parviflora AmSTrop Mediterranean, semi-arid DDHagri, DWH rare

97 Gamochaeta pensylvanica AmC, AmN Mediterranean DDHrdsid rare

98 Helianthus annuus AmN Mediterranean, semi-arid DDHagri, DDHrdsid, UE common

99 Heliotropium curassavicum AmN Mediterranean NWH, DWH very rare

100 Heterotheca subaxillaris AmNTemp Mediterranean NDH, DDHagri, DDHrdsid common

101 Ipomoea aquatica AsSE Mediterranean NWH very rare

102 Ipomoea cairica AfTrop Mediterranean DDHagri, UE common

211
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Scientific Name Origin Presence in Israel Habitat Occurrence

DDHagri, DDHrdsid, DDHrefsit,

103 Ipomoea hederacea AmC, AmSTrop Mediterranean very rare
DWH

104 Ipomoea lacunosa AmN Mediterranean DWH very rare

AmC, AmSTrop, AfTrop,

105 Ipomoea pes-caprae Mediterranean NDH very rare
AsTrop, AuN

106 Ipomoea triloba AmC Mediterranean DDHagri, DWH very rare

107 Lantana camara AmSTrop Mediterranean, desert NDH, DDHagri common

108 Lepidium ruderale EuCont, AsTempW Mediterranean DDHagri, DDHrdsid very rare

109 Lepidium sativum AsTempW Mediterranean DDHagri very rare

110 Leptochloa mucronata AmN, AmC, AmSTrop Mediterranean DDHagri very rare

111 Leptochloa uninervia AfTrop Mediterranean DWH very rare

112 Lupinus albus EuCont Mediterranean NDH very rare

113 Maireana brevifolia AuS semi-arid, desert DDHrdsid common

114 Matthiola incana EuMed Mediterranean UE very rare

115 Melia azedarach AuN, AsTrop Mediterranean DDHagri, DDHrdsid, UE common

116 Myriophyllum aquaticum AmS Mediterranean NWH, DWH very rare

117 Nicotiana glauca AmS Mediterranean, desert DDHall DWH, UE very common

118 Nothoscordum inodorum AmS Mediterranean DDHagri, UE very rare

119 Oenothera biennis AmN Mediterranean DDHrdsid very rare

 Scientific Name Origin Presence in Israel Habitat Occurrence

120 Oenothera drummondii AmC, AmN Mediterranean NDH, DDHrdsid common

121 Oenothera rosea AmC, AmNArid Mediterranean DDHagri very rare

122 Oenothera speciosa AmC, AmNArid Mediterranean DDHagri very rare

123 Opuntia ficus-indica AmC Mediterranean, semi-arid, desert NDH, DDHagri very common

NDH, DDHagri, DDHplfor,

124 Oxalis pes-caprae AfCape Mediterranean common
DDHrdsid, UE

125 Panicum antidotale AsTempC desert DDHagri, DWH very rare

126 Panicum capillare AmN Mediterranean, semi-arid DDHagri, DDHrdsid, DWH common

127 Panicum miliaceum AsTempC Mediterranean DDHagri very rare

128 Papaver somniferum EuMedE, AsTempW Mediterranean DDHrdsid very rare

NWH, DDHagri, DDHplfor,

129 Parkinsonia aculeata AmC, AmSTrop Mediterranean, semi-arid common
DDHrdsid, DWH

130 Parthenium hysterophorus AmC, AmSTrop Mediterranean DDHagri, DDHrdsid, DWH very rare

131 Paspalum dilatatum AmSTrop Mediterranean, semi-arid, desert NWH, DDHagri, DDHrdsid, DWH common

132 Paspalum distichum AmN, AmC, AmSTrop Mediterranean NWH, DWH rare

133 Pennisetum clandestinum AfTropE Mediterranean, semi-arid NWH, DWH rare

134 Pennisetum purpureum AfTrop Mediterranean NWH, DWH very rare

AsTempC, AsIndia,
135 Phyllanthus rotundifolius Mediterranean DDHagri rare
AsArP
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Scientific Name Origin Presence in Israel Habitat Occurrence

136 Physalis angulata AmN Mediterranean DDHagri, DDHrdsid rare

137 Physalis peruviana AmSTrop Mediterranean DDHagri very rare

138 Phytolacca americana AmN Mediterranean NWH, DWH rare

139 Pinus brutia EuMedE, AsTempW Mediterranean NDH, DDHagri, DDHplfor common

140 Pistia stratiotes AmSTrop Mediterranean NWH, DWH very rare

141 Populus alba Eu, AsTemp Mediterranean NWH very rare

142 Prosopis juliflora AmC semi-arid, desert NDH, DDHrdsid very rare

143 Ricinus communis AfTrop Mediterranean, semi-arid, desert NWH, DDHrdsid, DDHrefsit, DWH common

144 Robinia pseudoacacia AmNTemp Mediterranean DDHagri, DDHrdsid very rare

145 Salvinia molesta AmSTrop Mediterranean NWH very rare

146 Salvinia natans AmSTrop Mediterranean NWH very rare

NDH, DDHagri, DDHrdsid,

147 Schinus molle AmSTrop Mediterranean rare
DDHrefsit

148 Schinus terebinthifolius AmSTrop Mediterranean, semi-arid, desert NWH, DDHagri rare

149 Sesbania sesban AfTropE Mediterranean NWH, DWH very rare

150 Setaria pumila AfTrop & AsTrop (?) Mediterranean DDHagri, DDHrdsid very rare

151 Setaria verticillata Eu Mediterranean, semi-arid, desert DDHagri, DDHplfor, DDHrdsid rare

152 Sida acuta AmC Mediterranean DDHagri very rare

 Scientific Name Origin Presence in Israel Habitat Occurrence

153 Solanum cornutum AmNArid & AmC Mediterranean DDHagri very rare

DDHagri, DDHplfor, DDHrdsid,

154 Solanum elaeagnifolium AmNArid Mediterranean, semi-arid, desert common
DDHrefsit

155 Sorghum virgatum AfTropE (?) Mediterranean, semi-arid, desert DDHagri, DDHrdsid, DWH common

156 Sporobolus indicus AfTrop Mediterranean DDHagri rare

157 Tagetes minuta AmS Mediterranean DDHagri rare

158 Tetragonia tetragonoides AuN Mediterranean NDH, DDHrdsid rare
159 Tipuana tipu AmSTrop Mediterranean NDH very rare
160 Trianthema portulacastrum AfTrop, AsTrop Mediterranean DDHagri, DDHrdsid very rare
161 Verbesina encelioides AmNArid Mediterranean, semi-arid, desert NDH, DDHagri common
162 Veronica peregrina AmN Mediterranean NWH very rare
163 Vicia bengalensis EuMed, AfMed Mediterranean DDHagri very rare
164 Washingtonia robusta AmNArid Mediterranean NWH, DDHrdsid, DDHrefsit, UE common
NWH, DDHagri, DDHrdsid,
165 Xanthium spinosum AmSTrop Mediterranean, semi-arid, desert common
DDHrefsit, DWH, UE
Xanthium strumarium NWH, DDHagri, DDHrdsid,
166 AmN Mediterranean, semi-arid, desert common
complex DDHrefsit, DWH
215
 Jean-Marc Dufour-Dror (Ph.D)
Invasive
in Israel
Plants
Alien

Alien Invasive Plants in Israel Jean-Marc Dufour-Dror (Ph.D)