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Forestry Department

Food and Agriculture Organization of the United Nations

Forest Health & Biosecurity Working Papers

Alien Invasive Species: Impacts on Forests and Forestry

A Review

by
Beverly A. Moore

November 2005

Forest Resources Development Service Forest Resources Division Forestry Department

Working Paper FBS/8E FAO, Rome, Italy

Background

This paper is one of a series of FAO documents on forest-related health and biosecurity issues. The purpose of these papers is to provide early information on on-going activities and programmes, and to stimulate discussion.

The designations employed and the presentation of material in this publication do not imply the expression of any opinion whatsoever on the part of the Food and Agriculture Organization of the United Nations concerning the legal status of any country, territory, city or area or of its authorities, or concerning the delimitation of its frontiers or boundaries.
Further information on forest invasive species can be found at the FAO Alien invasive species Web site (www.fao.org/forestry/aliens) and information on forest health can be found at the FAO Forest health Web site (www.fao.org/forestry/site/pests).

Comments and feedback are welcome. For further information, please contact: Gillian Allard, Forestry Officer (Forest Protection and Health ) Forest Resources Division Forestry Department FAO, Viale delle Terme di Caracalla 00100 Rome, Italy Fax: + 39 06 570 55 137 Email: gillian.allard@fao.org Pierre Sigaud, Forestry Officer (Forest Genetic Resources ) Forest Resources Division Forestry Department FAO, Viale delle Terme di Caracalla 00100 Rome, Italy Fax: + 39 06 570 55 137 Email: pierre.sigaud@fao.org

FAO 2005

ALIEN INVASIVE SPECIES: IMPACTS ON FORESTS AND FORESTRY

A REVIEW

BY

BEVERLY A. MOORE CONSULTANT FOREST RESOURCES DEVELOPMENT SERVICE FORESTRY DEPARTMENT FAO NOVEMBER 2005

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ALIEN INVASIVE SPECIES: IMPACTS ON FORESTS AND FORESTRY TABLE OF CONTENTS

1. Introduction .........................................................................................................................................1 2. Alien invasive species affecting forests and forestry ...........................................................................2 3. Factors contributing to the introduction and spread of alien invasive species ..................................3 4. Impact of land use change and the forest sector on the introduction and spread of alien invasive species......................................................................................................................................................8 5. Positive impacts of introduced woody species on forests and forestry............................................. 11 6. Negative impacts of alien invasive species on forests and forestry .................................................. 13 7. Conflict species.................................................................................................................................. 17 8. Forest pest management options ...................................................................................................... 19 9. Conclusions........................................................................................................................................23 10. References ........................................................................................................................................24 Annex 1. Alien invasive species affecting forests and forestry ..............................................................34 Annex 2. Select alien invasive species affecting forests and forestry....................................................40 Annex 3. International and regional instruments and programmes.....................................................55

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1. Introduction

We are living in a period of the world's history when the mingling of thousands of kinds of organisms from different parts of the world is setting up terrific dislocations in nature Charles Elton, 1958. The ecology of invasions by animals and plants. An integral part of sustainable forest management includes measures to protect forests from natural threats such as fire, insects and diseases. Increasingly, an additional, and more severe threat, has been affecting the forest sector worldwide - alien invasive species. Alien invasive species are any species that are non-native to a particular ecosystem and whose introduction and spread causes, or are likely to cause, socio-cultural, economic or environmental harm or harm to human health. The increasing global movement of people and products, though beneficial to many people, is also facilitating the movement of alien species around world. These species may be unintentionally introduced to new environments in shipments of food, household goods, wood and wood products, new and used tires, animal and plant products, containers, pallets, internal packaging materials, and humans. In the absence of their natural predators, competitors and pathogens, they prosper in their new environments and spread at the expense of native species, affecting entire ecosystems. Not all invasive species have been inadvertently introduced, however. Particularly challenging to natural resource management are non-native species that have been intentionally introduced into an ecosystem to provide economic, environmental or social benefits. Many species of plants, trees and animals have been introduced outside their native ranges as ornamentals, for gardening, or for the pet industry, and these species have escaped to become serious problems in forests and other ecosystems. This is a considerable concern in the forest sector since many of the tree species used for agroforestry, commercial forestry and desertification control are alien or non-native to the area. It is vital to ensure that such species serve the purposes for which they were introduced and do not escape to cause negative effects on native ecosystems. While the definitions and impacts of invasive species on the forest sector are still debated and need reviewing in the context of forest management, a number of initiatives, programmes and activities have been initiated. Most programmes focus on damage caused to local forest ecosystems, or to particular species or group of species, by a given pest 1over a period of time. There is an overall lack of information on alien invasive species and the forest sector at the global scale. Information sharing is necessary in the planning and implementation of any strategy for the management of alien invasive species. This review concentrates, but not exclusively, on insects, diseases and woody species that have the potential to become invasive.

Any species, strain or biotype of plant, animal or pathogenic agent injurious to plants or plant products.

2. Alien invasive species affecting forests and forestry


Alien invasive species occur in all major taxonomic groups from micro-organisms to mammals. The Invasive Species Specialist Group (ISSG) of the Species Survival Commission (SSC) of the World Conservation Union (IUCN) has gathered information on 326 alien invasive species including 157 that negatively impact forests and the forest sector (www.issg.org/database [accessed on 23 August 2005]). Annex 1 provides a list of 204 species known to affect forests and the forest sector including the 157 species identified by ISSG. ISSG has also compiled a list of the One Hundred of the Worlds Worst Invasive Alien Species (www.issg.org/database/species/search.asp?st=100ss&fr=1&sts=#SpeciesList) which aims to collectively illustrate the range of impacts caused by biological invasion. Although incomplete, this list is a first attempt to rank the impact of alien invasive species. Included are 62 alien invasive species - four fungi, one flatworm, 10 insects, two molluscs, two amphibians, one reptile, two birds, 13 mammals, two grasses, six plants and 19 trees and shrubs - that impact forests and forestry. Annex 2 provides a more detailed discussion of these species.

3. Factors contributing to the introduction and spread of alien invasive species


Biological invasions are human-assisted humans intentionally and unintentionally introduce species into new areas or alter ecosystems in ways that promote invasions. Global factors, both primary and secondary, that support the introduction and spread of alien invasive species include: land use changes including forest sector activities (see Chapter 4); economics and trade; climate change and changes in atmospheric composition; tourism; conflict and reconstruction; regulatory regimes; biological control of pests; public health and environmental concerns. It is important to note here, that not all introduced species become invasive. It is widely accepted that only a small proportion of introduced species establish and only a small proportion of these species spread and become pests. This is often termed the tens rule whereby approximately 1 in 10 introduced species escapes to the wild, 1 in 10 of these introductions become established in the wild, and 1 in 10 of these established species becomes a pest (Vander Zanden, 2005). In addition, species that are known to be invasive elsewhere may not necessarily become invasive in a new environment. Economics and trade The openness of a countrys economy and the composition of its trade routes enhance the vulnerability of nations to biological invasions (Perrings et al., 2002; Taylor and Irwin, 2004). Invasions are also enhanced by the national importance of agriculture, forest and tourism sectors. A high importance generally leads to increases in the resources allocated to quarantine and protection however it also increases the opportunities for introduction and spread (FAO, 2001a). Dalmazzone (2000) investigated data on established alien species in 26 countries in Africa, Australia, Europe, and North and South America from the early 1960s to the early 1990s and found a clear correlation between economic variables and disturbances associated with human activities and a countrys vulnerability to biological invasions. The influence of such activities becomes more apparent when considering the problem of invasions in island ecosystems. With particularly vulnerable native biodiversity and a higher percentage of imports in comparison to continental areas, islands exhibit both economic and ecological reasons for the incidence of alien invasive species (Dalmazzone, 2000). In South Africa, its long colonial history, well developed infrastructure, and prosperous agriculture and forest sectors have contributed significantly to the introduction, establishment and spread of invasive alien plants. About 8 750 tree and plant species have been introduced into South Africa and of these, 161 species are considered highly invasive and many more are likely to become weeds in the future (van Wilgen et al., 2001). Globalization has led to more and faster trade, new travel and trading routes, and increased trade in livestock, pets, nursery stock, agricultural produce and forest products; all of which can facilitate the introduction and spread of alien invasive species (FAO, 2001a). Weed seeds, plant pathogens, larval or adult arthropods and other invertebrates, and even some vertebrate species can be transported on such commodities. Sand, gravel, coal and metal ores, and other inorganic commodities can also be contaminated with seeds, arthropods and pathogens (Cox, 1999). Unprocessed wood, wood products and nursery stock are also a major source of forest pests and diseases having introduced a number of devastating species into the United States such as chestnut blight (Cryphonectria parasitica), Dutch elm disease (Ophiostoma ulmi sensu lato), and white pine blister 3

rust (Cronartium ribicola) (OTA, 1993). It is believed that Pineus boerneri, a pine woolly adelgid, entered Kenya and Zimbabwe on scion material (Diekmann et al., 2002). In addition to the possibility of the commodities themselves carrying alien invasive species, the containers and vehicles in which they are transported can also facilitate invasions. Wood packaging material made of unprocessed raw wood, including pallets, crates, drums, skids, cases, and dunnage, can be a pathway for the introduction and spread of pests, in particular forest pests (McNeely et al., 2001; IPPC, 2002). The Asian longhorned beetle (Anoplophora glabripennis), emerald ash borer (Agrilus planipennis) and brown spruce longhorn beetle (Tetropium fuscum) are alleged to have arrived in North America among packing materials from Asia (Keiran and Allen, 2004). The Sirex woodwasp (Sirex noctilio) probably entered Argentina, Australia, New Zealand and South Africa in wooden packaging from Europe or North Africa and the red turpentine beetle (Dendroctonus valens) may have been introduced into Chinas pine forests through packaging made from infested North American wood (Keiran and Allen, 2004). The pinewood nematode (Bursaphelenchus xylophilus) the causative agent of pine wilt disease, has spread from its native North America to Asia and Europe in wooden packing materials (APHIS, 1999a; APHIS, 1999b). Containerized cargo can shelter alien species from microorganisms to reptiles and mammals and since inspecting such freight is very difficult and costly, many alien invasive species may enter a country undetected (OTA, 1993; Cox, 1999). No longer are the effects of alien species invasions being initially confined to areas around the ports of entry since containers are typically not unloaded until they reach their final inland destinations (OTA, 1993). Vehicles, including cars, trucks, trains, planes and ships, may also be contaminated with all types of pests and since the commercial and recreational movement of vehicles across international boundaries has increased, the threats are considerable (Cox, 1999). Railway sleepers have also been blamed for spreading many insect pests; it is suspected that Phoracantha semipunctata and P. recurva, both serious pests of eucalypts, entered South Africa in freshly-cut railway sleepers imported from Australia (EPPO, 2004). New trade activities, particularly in forest products, between China and the former Soviet Union and North America has led to a dramatic increase in pest and disease problems on both sides through accidental introduction of alien invasive species (McNeely et al., 2001; Normile, 2004). Climate change and changes in atmospheric concentration Human activities are releasing greenhouse gases such as carbon dioxide (CO2), methane, nitrous oxide, halocarbons and ozone into the atmosphere and the rising concentrations of such gases is changing climate (UNEP and UNFCCC, 2002). Global climate change has many environmental consequences including changes in species distributions and in their abundance within existing distributions as a result of direct physiological impacts on individual species and changes in abiotic factors, reproduction and recruitment opportunities, and interspecific interactions (McNeely et al., 2001). Climate change may produce more favourable conditions for alien invasive species. Once dominant species in native areas are no longer adapted to the environmental conditions of their habitat, it is likely that introduced species will displace them thus drastically changing successional patterns, ecosystem function and resource distribution (McNeely, 1999; Tilman and Lehman, 2001). For example, Oberhauser and Peterson (2003) investigated the possible effects of global climate change on distributions of migratory populations of monarch butterflies (Danaus plexippus) and concluded that predicted changes pose potentially significant threats to their survival. Climate, geography and other variables define the distribution limits of a species; however with changes in climate these limits are expanding, spreading species into higher latitudes and altitudes due to increased temperatures, humidity, precipitation and other climatic factors (McNeely, 1999; McNeely et al., 2001). Rouget 4

et al. (2002) noted that the current distribution of stands of invasive trees in South Africa was largely influenced by climatic factors. Warming trends may also allow for longer breeding seasons for invasive species, as observed in populations of the collared dove (Streptopelia decaocto) in Europe (Crooks and Soul, 1999). Since the development of insects is temperature dependent, it has been predicted that increasing temperatures will enhance the winter survival of insects and facilitate population increases and expansions in geographic range (Crooks and Soul, 1999; Mooney and Hofgaard, 1999). Changes in climate and environmental factors may also allow existing introduced species to become invasive (Mooney and Hofgaard, 1999). Climatically induced stress on plants and other species also reduces their ability to resist invaders and thus make them more vulnerable to insect or pathogen damage (Mooney and Hofgaard, 1999; McNeely et al., 2001). Climate change may alter production patterns and trade in agricultural and forestry commodities by species being grown more competitively in higher latitudes and altitudes. Since alien invasive species establish more easily in habitats disturbed by human and other factors, such changes can provide more opportunities for them to invade (McNeely et al., 2001). Climate change also affects the frequency and intensity of extreme climatic events which may have the greatest influence on alien invasive species by disturbing ecosystems and thus providing increased opportunities for dispersal and growth of invasive species (McNeely et al., 2001). In addition to the effects of climate change, increasing concentrations of greenhouse gases can have significant effects on the success of alien invasive species as well. Higher concentrations of carbon dioxide increase photosynthetic rates and water use efficiency of plants and ecosystems (Vitousek et al., 1997; Mooney and Hofgaard, 1999). The resulting increased soil moisture has potential to provide habitat for late-season annuals which may be invasive. Distributions of spruce budworm in Oregon have been observed to change in relation to increased atmospheric CO2 concentrations (Vitousek et al., 1997). However, increased levels of CO2 affect plant species differently which is likely to result in substantial changes in the species composition and dynamics of terrestrial ecosystems (Vitousek et al., 1997). A large component of the nitrogen cycle is atmospheric nitrogen which must be fixed before it can be used by most organisms. As a result, it is this fixed nitrogen that controls the productivity, carbon storage and species composition of many ecosystems (Vitousek et al., 1997). Changes in the deposition of nitrates from the atmosphere through the burning of fossil fuels, has resulted in large changes in vegetation, as observed in Western Europe, which may favour the growth of some alien invasive species (Mooney and Hofgaard, 1999). Conflict and reconstruction Conflict and civil unrest can contribute to the introduction and spread of alien invasive species in a variety of ways (FAO, 2001a). Civil unrest leads to the breakdown of phytosanitary and animal health controls and management systems, the loss of supply lines for materials as well as to the displacement of substantial numbers of people. Areas where there is civil unrest or war are more vulnerable to the entry of pests and diseases because of the lack of inspections and border controls and also because of the increased unregulated movement of military personnel and refugees. Displaced people and their belongings can be a dispersal mechanism for, or the source of, alien invasive species. Increased smuggling can relocate alien species to new regions. Inflows of food aid may be contaminated with pests and diseases. Difficulties in obtaining access to border areas because of landmines and other hazards make these areas difficult to survey. 5

Military transport, equipment and supplies, often covered with dirt or mud from the field, are effective means of dispersal for many species which can be detrimental to new environments. For example, the puncture vine (Tribulus terrestris), native to the Sahara Desert, may have been introduced into North America on the tires of military aircraft and vehicles returning from Europe after World War I (Cox, 1999). The brown tree snake (Boiga irregularis), native to New Guinea and neighbouring areas, was accidentally introduced to the island of Guam in the late 1940s or early 1950s most likely in military shipments of fruit (OTA, 1993; Cox, 1999). This snake is an arboreal, nocturnal predator of birds, their eggs and young that has nearly eliminated all native forest bird species. Black rats were introduced to the Midway Islands by navy ships during World War II where they drove many species of wildlife to extinction (Cox, 1999). A desert shrub, African rue (Peganum harmala) was apparently introduced inadvertently into New Mexico and Texas at World War II airfields (Cox, 1999). The agricultural pests, witchweed (Striga asiatica) and the golden nematode (Globodera rostochiensis) are also believed to have entered North America on returning military equipment (OTA, 1993; Cox, 1999). Ceratocystis fimbriata, a fungus that causes canker stain of plane trees (Platanus spp.) or platanus disease, was introduced from the United States to several Southern European ports at the end of the Second World War and subsequently spread through Italy and France (EPPO, undated). Heterobasidion annosum, a root rot of pine trees, was inadvertently introduced into Italy by American troops during World War II where it has resulted in an unprecedented mortality rate of stone pines (Pinus pinea) on the Presidential Estate of Castelporziano (Pilcher, 2004). It is believed that the pathogen was transferred in transport crates, pallets or other military equipment made from untreated lumber from infected trees. Emergency relief, reconstruction efforts, and humanitarian assistance after wars and disasters may also contribute to the introduction and spread of alien invasive species. Though little information is available, particularly in regards to pests of forests and forestry, foreign food aid has been accused of introducing agricultural pests into a number of African countries such as the larger grain borer (Prostephanus truncatus), unintentionally introduced into the United Republic of Tanzania in a food aid shipment in 1979 (FAO, 2001a). Regulatory regimes A countrys lack of regulatory regimes, including resources for prevention and enforcement measures (i.e. a sustainable institutional framework) as well as attitudes and views regarding risks, make it more vulnerable to invasions. Regulatory systems for managing alien invasive species are heavily dependent on the actions of both the government and private sectors and the effectiveness of such systems is determined by the level of resources that governments can provide as well as the technical capacity that exists nationally (FAO, 2001a). Countries vary in their tolerance of the risk of alien species. Since countries are linked to others by the transboundary movement of people, goods and services, the level of control applied by one country will in turn affect the vulnerability of other countries (Perrings et al., 2002). Regulatory systems can also break down, or when faced with new challenges in alien species control can become inadequate to respond effectively either because of systemic deficiencies or because safety measures are evaded (FAO, 2001a). Tourism With approximately 650 million tourists crossing international borders every year, the opportunities for the introduction and spread of alien invasive species is profound and increasing (McNeely et al., 2001). Travellers can intentionally transport living plant and animal species that can become invasive or they can carry fruits and other living or preserved plant materials that contain potentially invasive insects and diseases that can have profound effects on agriculture, forestry and other sectors (McNeely et al., 2001). Travellers themselves can also be the vectors for parasites and diseases between countries. Parks and protected areas have experienced increased biological invasions due to human activities (Mooney and Hofgaard, 1999). 6

Biological control of pests Another source of alien invasive species is the intentional importation and release of insects, snails, plant pathogens and nematodes for biological control of pests. Such species can escape into other unintended areas and become pests themselves. The United States Office of Technology Assessment (OTA) (1993) noted that of a total of 722 non-indigenous species imported for biological control, 237 species had become established in the United States, some of which have become detrimental. Public health and environmental concerns Concerns about the effects of pesticides on the environment and human health can also promote the spread of alien invasive species by allowing such species to spread unchecked (FAO, 2001a).

4. Impact of land use change and the forest sector on the introduction and spread of alien invasive species
Land use change has obvious direct effects on the worlds ecosystems through habitat destruction and alteration of competitive relationships but also indirect effects through alteration of fire frequency and nutrient and water balances (Mooney and Hofgaard, 1999). Land use change is a major contributor to the introduction and spread of alien invasive species which, in turn, are the second most important threat to biodiversity, behind habitat destruction. Activities of the forest sector can contribute to the introduction and spread of alien invasive species through forest utilization practices and the intentional introduction of species for commercial forestry, agroforestry and other purposes. Trade in forest products is also a contributor as was discussed in the section on trade in Chapter 3. Forest utilization, in particular practices such as timber harvesting, extraction of non-timber products, the construction of logging and transport roads and facilities for logging camps, and the conversion of natural forest to plantations, can have direct and indirect negative impacts on the ecological functions of forests and on forest biodiversity by promoting the invasion of alien species. For example, logging was found to be the prevailing factor influencing the establishment of alien plants areas in a national park in Madagascar and due to the dominance and persistence of the invasive species, the logged sites never recovered native species diversity (Brown and Gurevitch, 2004). As a result, the structure and diversity of the forest remained altered even long after logging operations has ceased. Clearcutting practices have also been shown to lead to the replacement of native ant communities with invasive nonindigenous species (Zettler et al., 2004). Forest roads provide essential access for timber extraction, management and monitoring of forest resources and as such are an important requirement for sustainable forest management and use. However when poorly designed and maintained, forest roads are often the cause of a variety of environmental problems associated with forest harvesting operations. Under some circumstances, forest roads may also initiate or accelerate the invasion of non-native species that ultimately displace native species. In addition, the increased levels of human activities in previously inaccessible areas, as facilitated by forest roads, cause many environmental problems including the possible introduction of alien species. Land use changes can promote the emergence or re-emergence of infectious diseases which degrade human health and that of other species (Morse, 1995; Vitousek et al., 1997; Bryan, 1999; Western, 2001). Infectious disease agents often, and perhaps typically, are alien invasive species since they are invaders over most of their range (McNeely et al., 2001). Activities of the forest sector contribute to the spread of infectious diseases in a variety of ways. Forest activities, such as clear-cutting and road building, may increase exposure of workers to infectious diseases such as human immunodeficiency virus (HIV), Ebola hemorrhagic fever, Marburg hemorrhagic fever, yellow fever, leishmaniasis, malaria and Ross River virus disease (Morse, 1995; Daszak, Cunningham and Hyatt, 2000; Chivian, 2002). The index case for the Ebola epidemic in the Democratic Republic of the Congo in 1995 is thought to be a charcoal maker that worked in the forest outside Kikwit (Sanchez et al., 1995). Logging can change the abundance, extent and quality of aquatic larval habitats for the Anopheles mosquito vectors of malaria and by disturbing the forest floor, creating depressions that catch and hold water, thus creating new sites for the development of more mosquitoes (Chivian, 2002). Deforestation can result in humans coming into closer contact with the vectors for such diseases as leishmaniasis, yellow fever, trypanosomiasis (both African sleeping sickness and Chagas disease), and 8

Kyasanur forest disease (Morse, 1995; Wilson, 1995; Chivian, 2001; Chivian, 2002; PAHO, 2004). Similarly, some animal reservoir hosts increase in abundance near the edges between forests and human settlements thus increasing the risk of human exposure to pathogens. The destruction of forest habitat may result in the removal, replacement or eradication of dominant vector species and sometimes, the replacement species are more effective vectors of disease as observed with loa loa (tropical eyeworm) and onchocerciasis (river blindness) (Chivian, 2002). Deforestation and desertification may also be accompanied by changes in the distribution of vectors such as ticks, blackflies, tsetse flies and Anopheles mosquitoes (Chivian, 2002). For example, forest degradation in the United States has led to the emergence of Lyme disease as a result of the loss of some predator and competitor species of white-footed mice which increased population densities of the mice in remnant forest fragments (Chivian, 2002). Higher population densities of the mice, which carry the bacteria (Borrelia burgdorferi) that cause Lyme disease, increase opportunities for ticks to acquire such infectious agents. Reafforestation activities can also affect the population dynamics of vectors and reservoirs which can promote the emergence of infectious diseases. Such activities in the United States and Europe have resulted in an increase in the population of deer and hence the deer tick, which is a vector of Lyme disease (Morse, 1995). The forest sector itself is a major source of alien invasive species. Many of the tree species used in commercial forestry in many parts of the world are alien or non-indigenous to the area. Hundreds of species have also been widely and successfully planted for a variety of purposes including afforestation, desertification and erosion control, and for the supply of fuelwood and other forest products. Such intentional introductions however, can bring about many unintended and costly consequences when these species escape cultivation and invade natural ecosystems. Alien tree plantations can have negative effects on the biodiversity and water resources of afforested areas (van Wilgen et al., 2001). Introduced species that spread from plantations to natural and semi-natural areas, and also into areas set aside for conservation and water production, have considerable impacts on ecosystem properties and functions (de Wit, Crookes and van Wilgen, 2001). In 2000, forest plantations covered 187 million hectares (FAO, 2000b). Pinus spp. (20 percent) and Eucalyptus spp. (10 percent) were the dominant species planted worldwide, followed by Hevea spp. (5 percent), Acacia spp. (4 percent), Tectona spp. (3 percent), other broadleaved species (18 percent), and other coniferous species (11 percent); the remaining percentage was unspecified. Globally, broadleaves made up 40 percent of forest plantation area with Eucalyptus the principal genus while coniferous species made up 31 percent of which Pinus was the principal genus; the remaining percentage was unspecified. Among the most widely planted trees, Acacia and Pinus species are the most prominent on weed lists and in reviews of invasive tree species (Richardson, 1999). This prominence is perhaps due to the fact that many species in both genera are highly capable of adapting to a wide variety of environments. Richardson and Rejmnek (2004), in a summary of data on conifer taxa from 40 countries, found that 80 alien taxa were known to be naturalized (13 percent of species) and 36 taxa were known invasives (6 percent). Twenty-eight of these known alien invasive conifer taxa belong to one family (Pinaceae) and 21 of these to one genus (Pinus). The Pinaceae has the highest proportion of invasive species than any other angiosperm family comprising mainly trees and shrubs. Considerable attention has been given to the effects of alien invasive trees from plantations in the southern hemisphere, in particular in South Africa. Plantations cover over 1.5 million ha of land area in South Africa and are an important part of the economy contributing US$300 million to the gross domestic product (GDP) and employing over 100 000 people (Richardson, 1998; FAO, 2000b). The forest sector has been one of the countrys major sources of alien species infestation; a large proportion (38 percent) of the area invaded by woody alien plants in South Africa is occupied by species used in commercial forestry, in particular Pinus and Acacia species (Richardson, 1998). Rouget et al. (2002) analyzed the distribution of forestry plantations and 9

invasive stands of Acacia mearnsii and Pinus spp. in South Africa and discovered that these two taxa account for 60 percent of the area under commercial plantations and 54 percent of the area invaded by alien trees and shrubs. Black wattle (Acacia mearnsii), introduced into South Africa from Australia in the mid-19th century, has been widely planted in the country and forms the basis of a small but significant industry. A highly invasive species, A. mearnsii has spread to an area of 2.5 million ha in South Africa where it negatively affects water resources, biodiversity, and the stability and integrity of riparian ecosystems (de Wit, Crookes and van Wilgen 2001). Due to their ability to fix atmospheric nitrogen, Australian Acacia species (A. cyclops and A. saligna), have radically altered nutrient-cycling regimes in nutrient poor fynbos ecosystems in South Africa (van Wilgen et al., 2001). The invasion of these species and some Pinus species into large areas of the tree-poor fynbos have caused many ecosystem level changes by altering factors such as biomass distribution, plant density and vegetation height, leaf-area index, litterfall and decomposition rates (van Wilgen et al., 2001). Such changes radically alter habitats resulting in major changes in the distributions of many animals. Invasive alien pines and acacias in South Africa can also reduce water tables (McNeely et al., 2001). Many countries are also reporting significant problems with introduced eucalypts from Australia which can be particularly harmful environmentally because their leaf litter contains chemical compounds that prevent other species from growing (McNeely et al., 2001). This can have significant consequences for the forest sector. Other notorious examples of invasive trees that negatively affect ecosystems include melaleuca and black cherry. The melaleuca or paper bark tree (Melaleuca quinquenervia), native to Australia, was introduced from South America into southern Florida in the United States as an ornamental in the early 1900s. Decades later, the population began to expand and the species is currently spreading at a rate of 11 000 ha a year throughout forests and grasslands of Floridas Everglades National Park where it damages native vegetation and wildlife (van Wilgen, Cowling and Burgers, 1996; Crooks and Soul, 1999; Pimentel et al., 2000; McNeely, 2004). The North American black cherry, Prunus serotina, was initially introduced into central Europe as an ornamental and over the years had been planted to produce valuable timber and to function as windbreaks, firebreaks and shelter for game (Starfinger et al., 2003). By the mid 1980s, the species was considered an aggressive invasive species, particularly in Germany, that formed dense thickets which threatened plant diversity and natural regrowth of indigenous forest tree species. Of the 2000 or so species used in agroforestry perhaps 10 percent are invasive (Richardson, 1999). While only one percent is considered highly invasive, these include many frequently planted species such as Casuarina glauca, Leucaena leucocephala and Pinus radiata (McNeely, 2004). The spread of introduced species from agroforestry plantings however are less of a threat than those from commercial forestry plantations since plantations are typically planted in much greater numbers over larger areas, afforested areas often adjoin natural vegetation, and spreading agroforestry trees will likely be held in check by agricultural practices and human use (Richardson, 1999). With the increase in afforestation and land use changes over the past few decades, the magnitude of such detrimental consequences has increased significantly (Richardson, 1998; de Wit, Crookes and van Wilgen, 2001). Generally, the species that cause the greatest problems are those that have been planted most widely and for the longest time and the areas most affected tend to have the longest histories of intensive planting (Richardson, 1998).

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5. Positive impacts of introduced woody species on forests and forestry


As mentioned in the previous chapter, the forest sector (and other sectors) often depends on introduced species to provide a variety of socio-economic, environmental and human health benefits to the forest sector and to the rural communities that depend upon forests. Many introduced species are highly regarded because of their exceptional adaptability to a wide variety of sites, their rapid growth, and the multiple uses of their products. With growing concerns about the degradation and loss of natural forests, planted forests and trees outside forests, composed most often of introduced woody species, are becoming increasingly more important sources of products such as timber, fibre and fuelwood. In developing countries, fuelwood is the prime source of energy, representing over 80 percent of the wood harvest (FAO, 1999). They also provide nonwood forest products, such as fruits, leaves, roots, honey, fibres, oils, resins, cosmetics and medicines, either from the planted trees themselves or from other elements of the ecosystem that they help to create. Such products contribute to the livelihoods of rural communities by providing food, medicine and employment as well as income from their sale. In countries with low forest cover, planted forests and trees outside forests constitute the main source of wood and non-wood forest products. In addition to providing valuable forest products, alien tree species planted in forest plantations and other areas help provide many vital ecosystem services such as: combating desertification; protecting soil and water; rehabilitating lands exhausted from other land uses; diversifying the rural landscape; maintaining biodiversity; enhancing carbon sequestration; providing amenity and shade. When planted in riparian areas, trees provide spawning beds for fish and molluscs and shade which aids in the reduction of eutrophication. Trees planted on farms help to increase soil fertility by providing organic matter through litter decomposition at the soil surface or through atmospheric nitrogen fixation (nitrogen-fixing trees), both of which contribute to improvements in food production. Along roads and highways, trees and plants not only add beauty to cities and towns but also provide shade and control outdoor noise and traffic pollution. Trees also play a major role in preserving the social and cultural values attached to forests, particularly as natural forests decrease in size through deforestation or are designated for other purposes. The forest sector also employs introduced species in biological control programmes to help combat pest problems with considerable success (McNeely, 1999). Practical reasons for using introduced species instead of native species in the forest sector include the following (Richardson, 1998; Richardson, 1999). Introduced trees often grow much faster than native species, particularly in the first years after planting. Native species are more difficult to manage silviculturally, in part because the biology of these species is often poorly known including information on how to collect, store and germinate seeds, how to produce seedlings in a nursery and how to manage them in a forest. Introduced species, on the other hand, have been well studied in a variety of settings. Seeds from introduced species are more readily available and easier to handle than seeds of native tree species. 11

In order to improve the trade balance by reducing the need for imported wood products, it is often necessary to develop local forest industries. Knowledge of markets and manufacturing technology is more readily available for introduced species such as pines and eucalypts. Positive impacts may be best achieved by careful selection of species and risk assessment prior to introduction and large-scale planting, coupled with management of introduced species in order to prevent them from becoming invasive.

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6. Negative impacts of alien invasive species on forests and forestry


Alien invasive species negatively impact the forest sector in economic, ecological and environmental, and social and health terms, though these impacts are almost never mutually exclusive. Economic Alien invasive species generate substantial costs to the forest sector in lost revenues, in expenses for their control and in lost conservation values and ecosystem services. Alien invasive species, in particular insect pests and diseases, can damage trees in all stages of development and affect the ability of both natural and planted forests to meet their management objectives (FAO, 2001b). The most direct economic impact of alien invasive species on the forest sector is related to the loss or reduced efficiency of production. Approximately US$4.2 billion in forest products are lost each year to alien insect pests and pathogens in the United States (Pimentel et al., 2000). In Canada, the damage resulting from past introductions of harmful invasive plant pests on agricultural crops and forestry has been estimated at CAD$7.5 billion annually and in the Canadian province of Manitoba alone, economic losses due to Dutch elm disease (Ophiostoma ulmi sensu lato) have been estimated at approximately CAD $30 million (Environment Canada, 2004). The detection of Asian longhorned beetle (Anoplophora glabripennis) in Canada poses a significant threat to both the hardwood products industry and the maple syrup industry, whose products were valued in 1997 at CAD$480 million and CAD$130 million respectively (Environment Canada, 2004). Such infestations of alien invasive species directly affect the quantities of forest products demanded or supplied thereby impacting global prices and markets (FAO, 2001a). Although quantitative estimates of the economic impacts are not readily available for other countries, alien invasive species no doubt significantly impact productivity. Possibly outweighing direct production costs, the introduction and spread of alien invasive species can have major implications for trade which will depend on the policy response of trading partners to news about outbreaks, the importance of the traded commodities, the extent of the damage, and the demand and supply elasticities (FAO, 2001a). In addition to these direct production and trade costs, the associated control costs, including the costs of inspections, monitoring, prevention, and response, of even just a few species can be enormous. The United States Forest Service currently spends approximately US$11 million annually on control of the gypsy moth (Lymantria dispar), a major pest of forest and ornamental trees (Pimentel et al., 2000). Removal of elm trees affected by Dutch elm disease (Ophiostoma ulmi sensu lato) costs approximately US$100 million per year (Pimentel et al., 2000). In South Africa, the cost of controlling alien plant invasions has been estimated to be approximately US$1 200 million although some have challenged this estimate (Nyoka, 2003). Ecological and environmental The full economic costs of invasions include not only the direct damage and control costs but also the effects on the ecosystems themselves. The ecological and environmental impacts of alien invasive species can be felt by all levels of organization including the gene, species, habitat and ecosystem level. Genes. If introduced or spread into habitats with closely related species, alien invasive species could interbreed with native species resulting in changes to the genetic makeup of either species (Secretariat of the Convention on Biological Diversity, 2003a). The possible negative consequences of such alterations include reduction in the survival of either species, creation of a more successful invader, or the creation of hybrids that could be more susceptible to certain pests and pathogens. Of recent concern to the forest sector is the 13

impact of possible introduction of new tree genotypes (non-local provenances or genetically improved planting stock) resulting in the creation of hybrids and the resulting loss of gene pools that may have acquired specific characteristics through local adaptation (FAO, 2005). The issue has not been extensively studied in forest trees, except possibly in the European black poplar (Populus nigra) (Cock, 2003). Species. Alien invasive species can influence species diversity, richness, composition and abundance. At the species level, direct effects of alien invasive species occur through processes such as the predation of, competition with, and pathogen and parasite transmission to individual organisms, eventually leading to population declines and species extinctions (Loehle, 2003; Secretariat of the Convention on Biological Diversity, 2003a). Through direct impacts on species or through alterations of habitats, alien invasive species are responsible for placing 762 forest species at risk (IUCN, 2005). The loss of such species is leading to a more homogenous world which is perhaps the biggest threat to global biological diversity, behind habitat loss (Perrings, Williamson and Dalmazzone, 2000; McNeely et al., 2001; Richardson and Rejmnek, 2004). Some examples of species level impacts in forests include the following. Miconia (Miconia calvescens), a tropical American tree introduced to French Polynesia in 1937, has significantly altered the forests of French Polynesia and other Pacific islands by shading out all native plants and promoting erosion and landslides with its shallow roots (Denslow, 2002). The invasive weeds, Scotch broom (Cytisus scoparius) and gorse (Ulex europaeus), are repressing the regrowth of the commercially important Douglas fir (Pseudotsuga menziesii) in British Columbia, Canada (FAO, 2000a). The Asian shrub, Amur honeysuckle (Lonicera maackii), has invaded many eastern North American areas where it impacts the natural regeneration of forests by reducing tree seedling density and growth and fecundity of perennial forest herbs (Gorchov and Trisel, 2003; Miller and Gorchov, 2004). Habitats. Through their impacts on species and ecosystem processes, alien invasive species can result in the fragmentation, destruction, alteration or complete replacement of habitats which in turn, has cascading effects on even more species and ecosystem processes (McNeely et al., 2001; Secretariat of the Convention on Biological Diversity, 2003a). Some examples of these impacts in forests include the following. Alien invasive diseases and pests have caused major changes in the composition of forests in eastern North America over the past century, including the decline of species like chestnut, elm and hemlock (McNeely et al., 2001). Kizlinski et al. (2002) investigated the direct impacts of an invasion of the hemlock woolly adelgid (Adelges tsugae) and the indirect impacts of the removal of the infested trees in hemlock-dominated forests and found profound changes in the structure, composition and ecosystem function. The loss of hemlock trees in these forests has also resulted in significant changes in the composition and distribution of bird populations (Tingley et al., 2002). When alien invasive insect species threaten native insect species, they can also have cascading effects on insectivorous birds and on plants that rely on insects for pollination or seed dispersal (McNeely et al., 2001). Ecosystems. The impacts of alien invasive species at the ecosystem level include changes to trophic structures, changes in the availability of resources such as water and nutrients, and changes in the disturbance regimes (McNeely et al., 2001; Secretariat of the Convention on Biological Diversity, 2003a). Some examples of the impacts of alien invasive species on forest ecosystems include the following. Invasive alien plants and trees have decreased water supplies for nearby communities and increased fire hazards in South Africa (McNeely et al., 2001; van Wilgen et al., 2001; Petit et al., 2004). Invasive grasses that are particularly fire-prone may lead to a permanent loss of forests (Mooney and Hofgaard, 1999). Australian Acacia species, such as A. cyclops and A. saligna, have radically altered nutrient cycling regimes in nutrient poor ecosystems due to their ability to fix atmospheric nitrogen (van Wilgen et al., 2001).

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Social and health The full costs of invasions also include the social and health impacts of alien invasive species on humans, in particular to the rural communities depending on forests. As a result of the negative impacts of alien invasive species on native forest biodiversity, a loss of food sources and traditional medicines may be experienced thereby compromising not only the health of local people but also the livelihoods of those dependent on the collection and sale of such items for income. For small-scale landowners, alien invasive species can also decrease the value of their land. Forest workers, as part of their jobs, and people living in and around forests are more exposed to alien invasive species such as the reservoirs and hosts of many emerging infectious diseases. Examples of such diseases include Lyme disease, human immunodeficiency virus (HIV), Ebola and Marburg hemorrhagic fevers, malaria, yellow fever, leishmaniasis, trypanosomiasis and Kyasanur forest disease (Morse, 1995; Sanchez et al., 1995; Wilson, 1995; Daszak, Cunningham and Hyatt, 2000; Chivian, 2001; Chivian, 2002; Cinco et al., 2004). Chapter 4 has a more detailed discussion of the connections between the forest sector, emerging infectious diseases and human health. People living in and around invaded forest areas may also suffer allergic or other negative reactions to the alien invasive species themselves or to the measures used to control them such as chemical and biological pesticides. A commonly planted tree for land restoration and as a source of forest products, mesquite (Prosopis juliflora) is a major cause of allergies in India, Kuwait, Mexico, Saudi Arabia, South Africa, the United Arab Emirates and the southwestern United States (Killian and McMichael, 2004). Sensitivity to mesquite pollen has been shown to result in asthma, rhinitis and conjunctivitis (Killian and McMichael, 2004). In Mongolia, children living close to areas infested with Dendrolimus sibiricus have experienced significant allergic reactions to the hairy caterpillars that have entered their homes. The hairs on larvae and egg masses of gypsy moth (Lymantria dispar) also cause allergies in some people (Allen, Miller and Tyler, 1991; ISSG Global Invasive Species Database, 2005). In the United States, forest workers working in areas heavily infested with the tussock moth caterpillar (Orgyia pseudotsugata) experienced itching of the skin and eyes, nasal discharge, cough and respiratory difficulty (Press et al., 1977). Estimating the costs of alien invasive species Estimates of the full costs of biological invasions are rare because of the difficulty in approximating the costs of a problem with so many components, many of which are difficult if not impossible to quantify such as the impacts of alien invasive species on biodiversity, ecosystem functions, human health and other indirect costs such as the impacts of control measures. No estimates of the costs to the forest sector on a global scale have been made. A few attempts at estimating the costs of alien invasive species have been made. Pimentel, Zuniga and Morrison (2005) estimates that the 50 000 alien species in the United States cost almost US$120 billion in environmental damages and losses yearly. Pimentel et al. (2000) gave an estimate of US$137 billion per year. Pimentel et al. (2001) looked at over 120 000 alien species of plants, animal and microbes that have invaded Australia, Brazil, India, South Africa, the United Kingdom and the United States causing significant economic losses in the agriculture and forest sectors and negatively affecting ecosystems. They estimated that the total cost in the six countries was US$314 billion in damages per year - Australia ($13 billion), Brazil ($50 billion), India ($116 billion), South Africa ($7 billion), the United Kingdom ($12 billion) and the United States ($116 billion). OTA (1993) concluded that about 4 500 exotic species occur in the United States and that about 20 percent of them have caused serious economic and environmental harm. The cumulative loss caused by 79 of these species was estimated at almost US$97 billion for the period 1906 to 1991.

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While these estimates do not take all components into account, they nonetheless illustrate the enormity of the costs of alien invasive species.

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7. Conflict species
While many introduced species are highly regarded because of the benefits they can provide, these same species have in some cases become serious threats to forests and the forest sector. Such conflict species are a considerable problem from a management perspective requiring a clear and unbiased analysis of the costs and benefits of their use. Some examples of conflict species include the following. Pinus and Eucalyptus species are the most important introduced species used in commercial forestry enterprises worldwide and most particularly in the tropics and subtropics (Richardson, 1998; Richardson and Higgins, 1998; FAO, 2000b). Many introduced commercial species such as rubber (Hevea brasiliensis), coconut (Cocos nucifera) and oil palm (Elaeis guineensi) are becoming important sources of wood and fibre (FAO, 2000b). Several of these alien forest trees have spread beyond the areas in which they were planted with devastating impacts. The main impacts are considered to be caused by reduced structural diversity, increased biomass, disruption of existing vegetation dynamics and altered nutrient cycling (Richardson, 1998; Richardson and Higgins, 1998). Many species of Australian Acacia have been introduced into the Cape Floristic Region of South Africa for timber, fuelwood and building materials (A. mearnsii); for tannins which are used by leather industries (A. saligna, A. mearnsii); and for sand stabilization (A. cyclops, A. saligna) (McNeely, 1999; de Wit, Crookes and van Wilgen, 2001; Matthews and Brand, 2004). Such species have radically altered habitats for wildlife resulting in major changes in the distribution of species, particularly birds, and nutrient cycling regimes in nutrient poor ecosystems due to their ability to fix atmospheric nitrogen (van Wilgen et al., 2001). They have also decreased water supplies for nearby communities and increased fire hazards (McNeely et al., 2001; van Wilgen et al., 2001; Petit et al., 2004). Leucaena leucocephala has been widely introduced as a source of timber, fuelwood, fodder and shade and is also used to restore degraded lands, improve soils and stabilize sands. Leucaena is a fast-growing, nitrogenfixing tree that is tolerant of arid conditions and saline soils and as such is highly regarded in arid regions in Asia and Africa (Matthews, 2004; Matthews and Brand, 2004). In areas where it has been introduced however, this species tends to form dense impenetrable thickets and readily invades forest margins, roadsides, wastelands, riparian areas and agricultural lands (McNeely, 1999). Also, the toxicity of its seeds and foliage decreases its value as a source of fodder (McNeely, 1999). Prosopis juliflora, introduced 70 years ago in the Thar Desert of India, has dense green vegetation which is very useful in controlling soil erosion, reducing the aridity of the area, and providing a source of fuelwood as well as fodder and shelter for both wild and domesticated animals (McNeely, 1999). Such benefits however are being overshadowed by the negative impacts of this species. P. juliflora displaces native flora resulting in reduced biodiversity and reduced diversity of products available to rural communities (McNeely, 1999). Its dense impenetrable thickets also render invaded lands useless for agricultural purposes (Richardson, 1998). Salt cedar (Tamarix spp.), introduced from central Asia to the southwestern United States nearly 200 years ago to control erosion along river banks, now forms dense thickets on more than 400 000 ha of riparian habitat severely impacting hydrological systems (McNeely et al., 2001). Australian brush-tailed possums (Trichosurus vulpecula), introduced into New Zealand for a successful fur industry, have caused considerable damage to native forests by changing forest composition and structure through the defoliation and eradication of preferred food plants (McNeely, 1999).

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The flatworm Platydemus manokwari has been introduced into many areas where it successfully controls populations of another alien invasive species, the giant African snail, Achatina fulica (McNeely, 1999). Although successful as a biological control agent, P. manokwari is now considered a significant threat to native gastropod species, including rare and endemic species, in the areas where it was introduced. It is vital to ensure that such species serve the purposes for which they were introduced and do not escape to cause negative consequences on native ecosystems.

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8. Forest pest management options


Two sets of complementary strategies for dealing with alien invasive species are: prevention and early detection; and response, which includes eradication (rarely achieved), containment, control and mitigation. Many agencies have been advocating the concept of biosecurity which is a strategic and integrated approach that encompasses the policy and regulatory frameworks (including instruments and activities) that analyse and manage risks in the sectors of food safety, animal life and health, and plant life and health, including associated environmental risk (FAO, 2003). It covers the introduction of insect pests and diseases, the introduction and release of genetically modified organisms and their products, and the introduction and management of alien invasive species and genotypes (FAO, 2003). Unless otherwise noted, the information presented in this section is taken from Wittenberg and Cock (2001). Prevention and early detection Prevention is the first line of defence against biological invasions and is also the most cost effective since once an alien invasive species becomes established, it is extremely difficult and hence costly to eradicate it. An important first step in prevention is identification of the species capable of becoming invasive, the possible susceptible sites and more importantly, the pathways in which they can be introduced. The more comprehensive approach of identifying pathways rather than individual species results in a greater concentration of effort where pests are more likely to enter a country which not only avoids wasting resources elsewhere but also helps in the identification of more species, vectors and pathways. Once pathways are identified then potential prevention tools and methods can be more specifically developed. The International Plant Protection Convention (IPPC) is an international treaty with a purpose to secure action to prevent the spread and introduction of pests of plants and plant products, and to promote appropriate measures for their control. This convention specifically governs transboundary movement of all plants and plant products (including forest products) and the scope is not limited to agricultural plants. It is governed by the Interim Commission on Phytosanitary Measures (ICPM) which adopts International Standards for Phytosanitary Measures (ISPMs). All of these ISPMs have direct relevance to the forest sector including guidelines for the export, shipment, import and release of biological control agents and other beneficial organisms (ISPM No. 3), guidelines on pest risk analysis (ISPM No. 2, 11 and 21), guidelines on pest eradication programmes (ISPM No. 9), guidelines on pest status and reporting (ISPM No. 8 and 17), and guidelines for regulating wood packaging materials in international trade (ISPM No. 15) (IPPC, 2005). Some of the important tools used to prevent the entry and establishment of alien invasive species include: public information and education; risk assessments and environmental impact assessments for intentional introductions; national and international regulations on prevention and quarantine measures and their enforcement with inspections and fees; treatment of imported commodities, including through fumigation, immersion, spraying, temperature treatment, ultraviolet sterilization, and pressure; technically justified trade restrictions; emergency measures or actions if a pest should be detected before establishment (McNeely et al., 2001). Early detection of alien species should be based on a system of regular surveys general, site-specific or species-specific to identify newly established species. Although not all alien species become invasive, the costs of those that do become invasive suggest that a precautionary approach to the issue is best. If alien species are identified early, the chances for eradication will be high in particular because for some invasive species there can be a long lag period between initial introduction and subsequent population explosion

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(Crooks and Soul, 1999; Parker, 2004). The longer species go undetected the fewer the options for its control or eradication and the more expensive any intervention will become. The IPPCs ISPM No. 5 provides the international standard to which general surveillance should be conducted for plants (IPPC, 2005). Early detection is highly dependent on the capacity of individuals to recognize both native and alien species (See Box 1). As a result, a large component of this step is training, not only of national professionals responsible for surveying but also for any persons that spends time in the natural environment such as farmers, gardeners, forest workers, ecologists, tourism workers, photographers and hikers. Trained professional national workers should be able to not only recognize native and alien species and the ecological effects of alien species but they also should be able to use databases, keys, manuals and other identification sources. Early warning systems which include lists and datasets of recorded or potentially invasive species, in given countries, time sets and conditions are important tools in this regard. Finally a contingency plan outlining the actions that should be taken once an alien invasive species has been identified or an invasion is suspected, should be developed. Box 1. Attributes of potentially invasive species and susceptible ecosystems Though much is known about the factors that contribute to the introduction and spread of alien invasive species, little is known about the attributes of alien invasive species and susceptible habitats. Isolated ecosystems, evolutionarily and geographically, are particularly vulnerable to biological invasions while deserts, semi-deserts, tropical dry forests and woodlands, and arctic systems appear to be the least susceptible (McNeely et al., 2001; Perrings et al., 2002). Ecosystems with low species diversity, especially if predators and competitors are absent, seem to be more susceptible than species-rich systems with well-established species interactions (McNeely et al., 2001; Perrings et al., 2002). However, such species-rich ecosystems may be at risk to a greater range of invaders because of their greater diversity of habitats (McNeely et al., 2001). Frequent disturbance, slow recovery rate and fragmentation of communities promote plant invasions (Rejmnek, 1999). Some ecological factors that may allow introduced species to spread include the following (Pimentel et al., 2000). Lack of predators, competitors and parasites. Ability of an alien parasite to switch to a new host. Ability to be an effective predator. Availability of artificial or disturbed habitats. High adaptability to novel ecosystems. Efficient dispersal. While the success of alien invasive species is often explained by simply placing an introduced species in a favourable environment, some have suggested that the lack of enemy pressure actually results in a reallocation of resources from defence, as it is no longer needed in enemy-free habitats, to growth and reproduction (Petit et al., 2004; Withgott, 2004). Others have suggested that this may not be the case for all circumstances and there is likely not a simple trade-off between defence mechanisms and growth (Withgott, 2004). Identification of the life-history traits of potentially invasive species has been attempted with some success, primarily for woody species. For conifer species, invasiveness is associated with: small mean seed mass (<50 mg); short juvenile period (<10 years); and short intervals between large seed crops (Richardson and Rejmnek, 2004). A short juvenile period and short intervals between large seed crops results in early and consistent reproduction. A short juvenile period may also be related to fast growth in general and small mean seed mass is associated with a greater number of seeds produced which are better dispersed, a high initial

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germinability and a shorter chilling period to overcome dormancy. Examples of alien invasive species with such life-history characteristics include Cryptomeria japonica, Larix deciduas, Picea sitchensis, Pinus contorta, Pinus strobus and Pseudotsuga menziesii. Another important consideration is the fact that some alien invasive species explode quickly while others may have a long lag period between initial introduction and subsequent population growth and expansion. Some reasons for such time lags include the following (Crooks and Soul, 1999). The area in which the species has been introduced may become more susceptible to invasion. The species may have been undergoing rapid yet undetected expansion and some environmental change may trigger invasion. Several decades may be needed to build up large enough populations to have significant reproductive potential. Newly introduced species may be confined to restricted habitats until mutations favourable for further colonization became available. Knowing more about which species invade and which ecosystems are more susceptible will improve and refine capacities to prevent, or detect in a timely manner, biological invasions. Response When the preventative and early detection measures have failed to stop the introduction of alien invasive species, eradication is the preferred next method of action. Eradication involves the elimination of the entire population of an alien species, including resting stages, in managed areas. As a rapid response to early detection of an alien species, eradication is often the key to a successful and cost-effective solution. Careful analysis of the costs involved and the likelihood of success must be made before any eradication attempts are made. Some groups of organisms are more suitable for eradication such as plants, terrestrial vertebrates and some terrestrial invertebrates and in some cases insects. Also, well established populations and large areas of infestation are unsuitable for such programmes. Successful eradication programmes must: be scientifically based; ensure that all individuals of the target population are susceptible to the technique being used thus ensuring that eradication of all individuals is achievable; build support from the public and all relevant stakeholders; ensure that the legal and institutional framework is sufficient for dealing with the issue; secure sufficient funding; ensure through prevention measures that there is no immigration of the target species into the area; put in place a method to detect the last survivors; include a subsequent monitoring phase to ensure that eradication has been achieved, and to prevent reinvasion; ensure that techniques are environmentally, socially and ethically acceptable; include any necessary measures to restore ecosystems after eradication. Eradication programmes involve several control methods as outlined in the control section below. The IPPCs ISPM No. 09 specifically provides general guidance for eradication programmes targeting plant pests and alien invasive species (IPPC, 2005). Containment is a special form of control aimed at restricting the spread of an alien invasive species and to contain the population in a defined geographical range. The methods used are the same as those described for prevention, eradication and control. An important component of containment programmes is the ability to rapidly detect new infestations of the alien species spreading from the defined containment area or into new

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areas, so that control measures can be implemented quickly. In cases where eradication is not possible, containment of the alien invasive species into a defined area can be very effective in saving other regions of a country. The long-term reduction in density and abundance of an alien invasive species to below a pre-set acceptable threshold is the aim of control programmes. Suppression of invasive populations below such thresholds can favour native species. All control programmes need long-term funding and commitment. Control methods, including mechanical control, chemical control, biological control, habitat management, and hunting or a combination of these, have all been used successfully in controlling alien invasive species. Not all options for the control of alien invasive species are practical, effective, economically justifiable, or environmentally sound for application in forests. In the forest sector, control measures should be integrated to maximize yield and profit while minimizing negative environmental impacts. Some other options that are suitable include cultural techniques, such as seedling management, planting patterns, buffer zones, ecoclimatic matching, and pest and pathogen resistance; mechanical methods, such as cutting, bulldozing and shading; biological control; and integrated pest management. If eradication, containment and control methods have failed in managing an alien invasive species, the only option is to accept the species while mitigating the impacts to other species and the environment. The focus here is on protecting native species rather than harming the alien invasive species. Mitigation methods can include translocation of a population of species to an ecosystem that has not been invaded and alterations in the behaviour of desired species such as conditioning species to use breeding areas inaccessible to the invaders or artificial feeding sites. Communication Central to any effective alien invasive species management programme is rapid and transparent communication with all stakeholders which helps ensure maximum stakeholder participation and increased awareness of the issues. Such communication is vital during the risk assessment, prevention and management phases. For plants and plant products, governments have specific information exchange obligations in this regard (IPPC, 2005) to create awareness in neighbouring countries and among trading partners. Such communication can greatly facilitate trade by increasing transparency and trust between trading partners.

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9. Conclusions
Alien invasive species impact, and will continue to impact forests and the forest sector. The impact will be felt not only at the biological level, through technical responses to invasiveness, but also at policy, regulatory and trade levels. While the forest sector has long been considered a victim, there is growing evidence of and attention to its contribution to invasiveness. A major part of the solution to the problem of alien invasive species will come from local and national actions with regards to early warning systems, eradication and control attempts as well as increased awareness and political pressure (Mooney and Hobbs, 2000). The other part of the solution requires bilateral, regional and international efforts, since the issue of alien invasive species is global in scope. Numerous international and regional programmes and instruments, binding and nonbinding, have been developed to address the problem of alien invasive species, some with direct or indirect implications for forests and the forest sector (Annex 3). In order to reduce the impacts of alien invasive species on the forest sector, in particular, it has been recommended that the issue be better integrated into existing sustainable forestry frameworks such as the Montral Process and forest certification programmes (Chornesky et al., 2005). An interdisciplinary approach is needed to fully understand, prevent and reduce the harmful effects of alien invasive species one that incorporates biological, ecological and social sciences, economics, policy analysis, decision sciences, informatics and engineering (Mooney and Hobbs, 2000; FAO, 2003; Chornesky et al., 2005). Clearly, alien invasive species are serious threats to forests and the forest sector. It is also clear that introduced species have many positive impacts on the forest sector and to sustainable development in general. Such conflicts of interest require a balanced analysis of the costs and benefits of the introduction of alien species which will help strengthen policy and management decisions, in particular those dealing with international trade and pest suppression efforts (de Wit, Crookes and van Wilgen 2001; Chornesky et al., 2005).

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10. References
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Annex 1. Alien invasive species affecting forests and forestry


ORGANISM
TYPE

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30

SPECIES Brenneria (Erwinia) quercina Xylella fastidiosa Batrachochytrium dendrobatidis Ceratocystis fimbriata Cronartium ribicola Cryphonectria parasitica Discula destructiva Ophiostoma ulmi sensu lato Phytophthora cinnamomi Phytophthora ramorum Sirococcus clavigignenti-juglandacearum Platydemus manokwari Varroa destructor Adelges piceae Adelges tsugae Aedes albopictus Agrilus planipennis Ambrosiodmus lewisi Anoplolepis gracilipes Anoplophora chinensis Anoplophora glabripennis Anthonomus grandis Apis mellifera scutellata Aulacaspis yasumatsui Batrachedra pinicolella Callidiellum rufipenne Callidiellum villosulum Cinara cupressivora Coptotermes formosanus Corythucha arcuata

Bacteria Bacteria Fungus Fungus Fungus Fungus Fungus Fungus Fungus Fungus Fungus Flatworm Arachnid Insect Insect Insect Insect Insect Insect Insect Insect Insect Insect Insect Insect Insect Insect Insect Insect Insect

REFERENCE(S) www.eppo.org/QUARANTINE/Alert_List/bacteria/brenneria.htm www.issg.org/database/species/ecology.asp?si=326&fr=1&sts= www.issg.org/database/species/ecology.asp?si=123&fr=1&sts= www.eppo.org/QUARANTINE/fungi/Ceratocystis_fimbriata_platani/CERAFP_ds.pdf www.issg.org/database/species/ecology.asp?si=550&fr=1&sts= www.issg.org/database/species/ecology.asp?si=124&fr=1&sts= www.issg.org/database/species/ecology.asp?si=224&fr=1&sts= www.issg.org/database/species/ecology.asp?si=130&fr=1&sts= www.issg.org/database/species/ecology.asp?si=143&fr=1&sts= www.issg.org/database/species/ecology.asp?si=563&fr=1&sts= www.issg.org/database/species/ecology.asp?si=711&fr=1&sts= www.issg.org/database/species/ecology.asp?si=133&fr=1&sts= www.issg.org/database/species/ecology.asp?si=478&fr=1&sts= www.issg.org/database/species/ecology.asp?si=226&fr=1&sts= www.issg.org/database/species/ecology.asp?si=230&fr=1&sts= www.issg.org/database/species/ecology.asp?si=109&fr=1&sts= www.issg.org/database/species/ecology.asp?si=722&fr=1&sts= www.ncrs.fs.fed.us/4501/focus/default.asp www.issg.org/database/species/ecology.asp?si=110&fr=1&sts= www.eppo.org/QUARANTINE/insects/Anoplophora_malasiaca/ANOLMA_ds.pdf www.issg.org/database/species/ecology.asp?si=111&fr=1&sts= www.issg.org/database/species/ecology.asp?si=549&fr=1&sts= www.issg.org/database/species/ecology.asp?si=325&fr=1&sts= www.issg.org/database/species/ecology.asp?si=814&fr=1&sts= www.pestalert.org/pestnews.cfm#343%20 www.ncrs.fs.fed.us/4501/focus/imminent_threats/ www.ncrs.fs.fed.us/4501/focus/imminent_threats/ www.issg.org/database/species/ecology.asp?si=121&fr=1&sts= www.issg.org/database/species/ecology.asp?si=61&fr=1&sts= www.eppo.org/QUARANTINE/Alert_List/insects/crthar.htm 34

31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66

Ctenarytaina spatulata Diocalandra frumenti Euwallacea fornicatus Glycaspis brimblecombei Harmonia axyridis Hylastes opacus Hylurgops palliatus Hylurgus ligniperda Hyphantria cunea Linepithema humile Lymantria dispar Orgyia thyellina Orthotomicus erosus Oryctes rhinoceros Paratrechina longicornis Paysandisia archon Pheidole megacephala Phoracantha recurva Pityogenes bidentatus Platypus mutatus Rhynchophorus ferrugineus Rhynchophorus palmarum Scolytus schevyrewi Sirex noctillo Solenopsis geminata Solenopsis invicta Teia anartoides Tetropium fuscum Tomicus piniperda Uraba lugens Vespula pensylvanica Vespula vulgaris Wasmannia auropunctata Xyleborinus alni Xyleborus atratus Xyleborus californicus

Insect Insect Insect Insect Insect Insect Insect Insect Insect Insect Insect Insect Insect Insect Insect Insect Insect Insect Insect Insect Insect Insect Insect Insect Insect Insect Insect Insect Insect Insect Insect Insect Insect Insect Insect Insect

www.eppo.org/QUARANTINE/Alert_List/insects/ctenarytaina.htm www.eppo.org/QUARANTINE/Alert_List/insects/diocalandra.htm www.ncrs.fs.fed.us/4501/focus/default.asp www.eppo.org/QUARANTINE/Alert_List/insects/glybrim.htm www.issg.org/database/species/ecology.asp?si=668&fr=1&sts= www.ncrs.fs.fed.us/4501/focus/default.asp www.ncrs.fs.fed.us/4501/focus/default.asp www.ncrs.fs.fed.us/4501/focus/default.asp www.biosecurity.govt.nz/pests-diseases/forests/fall-webworm/index.htm www.issg.org/database/species/ecology.asp?si=127&fr=1&sts= www.issg.org/database/species/ecology.asp?si=96&fr=1&sts= www.biosecurity.govt.nz/pests-diseases/forests/white-spotted-tussock-moth/index.htm www.issg.org/database/species/ecology.asp?si=787&fr=1&sts= www.issg.org/database/species/ecology.asp?si=173&fr=1&sts= www.biosecurity.govt.nz/pests-diseases/animals/invasive-ants/crazy-ants/index.htm www.eppo.org/QUARANTINE/Alert_List/insects/paysar.htm www.issg.org/database/species/ecology.asp?si=132&fr=1&sts= www.eppo.org/QUARANTINE/Alert_List/insects/phoracantha.htm www.ncrs.fs.fed.us/4501/focus/default.asp www.eppo.org/QUARANTINE/Alert_List/insects/platypus.htm www.eppo.org/QUARANTINE/Alert_List/insects/rhycfe.htm www.eppo.org/QUARANTINE/Alert_List/insects/rhycpa.htm spfnic.fs.fed.us/exfor/data/pestreports.cfm?pestidval=163&langdisplay=english treesearch.fs.fed.us/pubs/viewpub.jsp?index=12997 www.issg.org/database/species/ecology.asp?si=169&fr=1&sts= www.issg.org/database/species/ecology.asp?si=77&fr=1&sts= www.biosecurity.govt.nz/pests-diseases/forests/painted-apple-moth/index.htm www.atl.cfs.nrcan.gc.ca/index-e/what-e/science-e/entomology-e/bslb-e/efpa-e.html www.ncrs.fs.fed.us/4501/focus/default.asp www.biosecurity.govt.nz/pests-diseases/forests/gum-leaf-skeletoniser/index.htm www.issg.org/database/species/ecology.asp?si=174&fr=1&sts= www.issg.org/database/species/ecology.asp?si=67&fr=1&sts= www.issg.org/database/species/ecology.asp?si=58&fr=1&sts= www.ncrs.fs.fed.us/4501/focus/default.asp www.ncrs.fs.fed.us/4501/focus/default.asp www.ncrs.fs.fed.us/4501/focus/default.asp

35

67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 98 99 100 101 102

Xyleborus glabratus Xyleborus pelliculosus Xyleborus pfeili Xyleborus similis Xylosandrus compactus Xylosandrus mutilatus Bursaphelenchus xylophilus Achatina fulica Euglandina rosea Bufo marinus Eleutherodactylus coqui Boiga irregularis Carpodacus mexicanus Molothrus ater Pycnonotus cafer Sturnus vulgaris Canis lupus Capra hircus Cervus elaphus Erinaceus europaeus Felis catus Herpestes javanicus Macaca fascicularis Mus musculus Mustela erminea Oryctolagus cuniculus Ovis aries Rattus exulans Rattus norvegicus Rattus rattus Sciurus carolinensis Suncus murinus Sus scrofa Trichosurus vulpecula Vulpes vulpes Arundo donax

Insect Insect Insect Insect Insect Insect Nematode Mollusc Mollusc Amphibian Amphibian Reptile Bird Bird Bird Bird Mammal Mammal Mammal Mammal Mammal Mammal Mammal Mammal Mammal Mammal Mammal Mammal Mammal Mammal Mammal Mammal Mammal Mammal Mammal Grass

www.ncrs.fs.fed.us/4501/focus/default.asp www.ncrs.fs.fed.us/4501/focus/default.asp www.ncrs.fs.fed.us/4501/focus/default.asp www.ncrs.fs.fed.us/4501/focus/default.asp www.issg.org/database/species/ecology.asp?si=175&fr=1&sts= www.ncrs.fs.fed.us/4501/focus/default.asp www.issg.org/database/species/ecology.asp?si=769&fr=1&sts= www.issg.org/database/species/ecology.asp?si=64&fr=1&sts= www.issg.org/database/species/ecology.asp?si=92&fr=1&sts= www.issg.org/database/species/ecology.asp?si=113&fr=1&sts= www.issg.org/database/species/ecology.asp?si=105&fr=1&sts= www.issg.org/database/species/ecology.asp?si=54&fr=1&sts= www.issg.org/database/species/ecology.asp?si=485&fr=1&sts= www.issg.org/database/species/ecology.asp?si=310&fr=1&sts= www.issg.org/database/species/ecology.asp?si=138&fr=1&sts= www.issg.org/database/species/ecology.asp?si=74&fr=1&sts= www.issg.org/database/species/ecology.asp?si=146&fr=1&sts= www.issg.org/database/species/ecology.asp?si=40&fr=1&sts= www.issg.org/database/species/ecology.asp?si=119&fr=1&sts= www.issg.org/database/species/ecology.asp?si=176&fr=1&sts= www.issg.org/database/species/ecology.asp?si=24&fr=1&sts= www.issg.org/database/species/ecology.asp?si=86&fr=1&sts= www.issg.org/database/species/ecology.asp?si=139&fr=1&sts= www.issg.org/database/species/ecology.asp?si=97&fr=1&sts= www.issg.org/database/species/ecology.asp?si=98&fr=1&sts= www.issg.org/database/species/ecology.asp?si=18&fr=1&sts= www.issg.org/database/species/ecology.asp?si=843&fr=1&sts= www.issg.org/database/species/ecology.asp?si=170&fr=1&sts= www.issg.org/database/species/ecology.asp?si=159&fr=1&sts= www.issg.org/database/species/ecology.asp?si=19&fr=1&sts= www.issg.org/database/species/ecology.asp?si=65&fr=1&sts= www.issg.org/database/species/ecology.asp?si=162&fr=1&sts= www.issg.org/database/species/ecology.asp?si=73&fr=1&sts= www.issg.org/database/species/ecology.asp?si=48&fr=1&sts= www.issg.org/database/species/ecology.asp?si=66&fr=1&sts= www.issg.org/database/species/ecology.asp?si=112&fr=1&sts=

36

103 104 105 106 107 108 109 110 111 112 113 114 115 116 117 118 119 120 121 122 123 124 125 126 127 128 129 130 131 132 133 134 135 136 137 138

Imperata cylindrica Microstegium vimineum Nassella neesiana Neyraudia reynaudiana Pennisetum clandestinum Abelmoschus moschatus Alliaria petiolata Andropogon virginicus Anredera cordifolia Celastrus orbiculatus Centaurea melitensis Chromolaena odorata Clematis vitalba Coccinia grandis Cryptostegia grandiflora Cytisus scoparius Dioscorea oppositifolia Dipogon lignosus Euonymus fortunei Euphorbia esula Gunnera tinctoria Hakea sericea Hedera helix Hedychium flavescens Hedychium gardnerianum Lespedeza cuneata Lonicera japonica Lonicera maackii Lythrum salicaria Merremia peltata Mikania micrantha Paederia foetida Passiflora tarminiana Polygonum perfoliatum Pueraria montana var. lobata Ranunculus ficaria

Grass Grass Grass Grass Grass Plant Plant Plant Plant Plant Plant Plant Plant Plant Plant Plant Plant Plant Plant Plant Plant Plant Plant Plant Plant Plant Plant Plant Plant Plant Plant Plant Plant Plant Plant Plant

www.issg.org/database/species/ecology.asp?si=16&fr=1&sts= www.issg.org/database/species/ecology.asp?si=686&fr=1&sts= www.issg.org/database/species/ecology.asp?si=458&fr=1&sts= www.issg.org/database/species/ecology.asp?si=303&fr=1&sts= www.issg.org/database/species/ecology.asp?si=183&fr=1&sts= www.issg.org/database/species/ecology.asp?si=15&fr=1&sts= www.issg.org/database/species/ecology.asp?si=406&fr=1&sts= www.issg.org/database/species/ecology.asp?si=200&fr=1&sts= www.issg.org/database/species/ecology.asp?si=776&fr=1&sts= www.issg.org/database/species/ecology.asp?si=156&fr=1&sts= www.issg.org/database/species/ecology.asp?si=264&fr=1&sts= www.issg.org/database/species/ecology.asp?si=47&fr=1&sts= www.issg.org/database/species/ecology.asp?si=157&fr=1&sts= www.issg.org/database/species/ecology.asp?si=348&fr=1&sts= www.issg.org/database/species/ecology.asp?si=347&fr=1&sts= FAO, 2000a; www.pfc.forestry.ca/biodiversity/broom_e.html www.issg.org/database/species/ecology.asp?si=296&fr=1&sts= www.issg.org/database/species/ecology.asp?si=804&fr=1&sts= www.issg.org/database/species/ecology.asp?si=575&fr=1&sts= www.issg.org/database/species/ecology.asp?si=83&fr=1&sts= www.issg.org/database/species/ecology.asp?si=836&fr=1&sts= van Wilgen et al., 2001 www.issg.org/database/species/ecology.asp?si=469&fr=1&sts= www.issg.org/database/species/ecology.asp?si=196&fr=1&sts= www.issg.org/database/species/ecology.asp?si=57&fr=1&sts= www.issg.org/database/species/ecology.asp?si=270&fr=1&sts= www.issg.org/database/species/ecology.asp?si=158&fr=1&sts= Miller and Gorchov, 2004; www.invasive.org/browse/subject.cfm?sub=3040 FAO, 2000a; www.invasive.org/browse/subject.cfm?sub=3047 www.issg.org/database/species/ecology.asp?si=163&fr=1&sts= www.issg.org/database/species/ecology.asp?si=42&fr=1&sts= www.issg.org/database/species/ecology.asp?si=632&fr=1&sts= www.issg.org/database/species/ecology.asp?si=336&fr=1&sts= www.issg.org/database/species/ecology.asp?si=582&fr=1&sts= www.issg.org/database/species/ecology.asp?si=81&fr=1&sts= www.issg.org/database/species/ecology.asp?si=844&fr=1&sts=

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139 140 141 142 143 144 145 146 147 148 149 150 151 152 153 154 155 156 157 158 159 160 161 162 163 164 165 166 167 168 169 170 171 172 173 174

Sphagneticola trilobata Tradescantia fluminensis Tradescantia spathacea Tussilago farfara Vinca major Wisteria floribunda Wisteria sinensis Elaeis guineensis Acacia mearnsii Acacia melanoxylon Adenanthera pavonina Ailanthus altissima Ardisia elliptica Berberis thunbergii Buddleja davidii Caesalpinia decapetala Cecropia peltata Cecropia schreberiana Cinchona pubescens Cinnamomum camphora Clidemia hirta Colubrina asiatica Dichrostachys cinerea Elaeagnus umbellata Euonymus alata Ficus rubiginosa Frangula alnus Hiptage benghalensis Kunzea ericoides Lantana camara Leucaena leucocephala Ligustrum robustum Ligustrum sinense Melaleuca quinquenervia Melia azedarach Miconia calvescens

Plant Plant Plant Plant Plant Plant Plant Palm Tree/Shrub Tree/Shrub Tree/Shrub Tree/Shrub Tree/Shrub Tree/Shrub Tree/Shrub Tree/Shrub Tree/Shrub Tree/Shrub Tree/Shrub Tree/Shrub Tree/Shrub Tree/Shrub Tree/Shrub Tree/Shrub Tree/Shrub Tree/Shrub Tree/Shrub Tree/Shrub Tree/Shrub Tree/Shrub Tree/Shrub Tree/Shrub Tree/Shrub Tree/Shrub Tree/Shrub Tree/Shrub

www.issg.org/database/species/ecology.asp?si=44&fr=1&sts= www.issg.org/database/species/ecology.asp?si=497&fr=1&sts= www.issg.org/database/species/ecology.asp?si=493&fr=1&sts= www.issg.org/database/species/ecology.asp?si=426&fr=1&sts= www.issg.org/database/species/ecology.asp?si=487&fr=1&sts= www.issg.org/database/species/ecology.asp?si=286&fr=1&sts= www.issg.org/database/species/ecology.asp?si=287&fr=1&sts= www.issg.org/database/species/ecology.asp?si=377&fr=1&sts= www.issg.org/database/species/ecology.asp?si=51&fr=1&sts= www.issg.org/database/species/ecology.asp?si=197&fr=1&sts= www.issg.org/database/species/ecology.asp?si=201&fr=1&sts= www.issg.org/database/species/ecology.asp?si=319&fr=1&sts= www.issg.org/database/species/ecology.asp?si=52&fr=1&sts= www.issg.org/database/species/ecology.asp?si=592&fr=1&sts= www.issg.org/database/species/ecology.asp?si=650&fr=1&sts= www.issg.org/database/species/ecology.asp?si=510&fr=1&sts= www.issg.org/database/species/ecology.asp?si=116&fr=1&sts= www.issg.org/database/species/ecology.asp?si=117&fr=1&sts= www.issg.org/database/species/ecology.asp?si=63&fr=1&sts= www.issg.org/database/species/ecology.asp?si=291&fr=1&sts= www.issg.org/database/species/ecology.asp?si=53&fr=1&sts= www.issg.org/database/species/ecology.asp?si=371&fr=1&sts= www.issg.org/database/species/ecology.asp?si=161&fr=1&sts= www.issg.org/database/species/ecology.asp?si=262&fr=1&sts= www.issg.org/database/species/ecology.asp?si=574&fr=1&sts= www.issg.org/database/species/ecology.asp?si=805&fr=1&sts= www.issg.org/database/species/ecology.asp?si=810&fr=1&sts= www.issg.org/database/species/ecology.asp?si=87&fr=1&sts= www.issg.org/database/species/ecology.asp?si=160&fr=1&sts= www.issg.org/database/species/ecology.asp?si=56&fr=1&sts= www.issg.org/database/species/ecology.asp?si=23&fr=1&sts= www.issg.org/database/species/ecology.asp?si=90&fr=1&sts= www.issg.org/database/species/ecology.asp?si=241&fr=1&sts= www.issg.org/database/species/ecology.asp?si=45&fr=1&sts= www.issg.org/database/species/ecology.asp?si=636&fr=1&sts= www.issg.org/database/species/ecology.asp?si=2&fr=1&sts=

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175 176 177 178 179 180 181 182 183 184 185 186 187 188 189 190 191 192 193 194 195 196 197 198 199 200 201 202 203 204

Mimosa pigra Morella faya Opuntia ficus-indica Paulownia tomentosa Pereskia aculeata Pinus pinaster Piper aduncum Pittosporum undulatum Populus alba Prunus serotina Psidium cattleianum Psidium guajava Rhamnus cathartica Rhodomyrtus tomentosa Robinia pseudoacacia Rosa multiflora Rubus ellipticus Rubus moluccanus Rubus rosifolius Schinus terebinthifolius Sesbania punicea Solanum mauritianum Spathodea campanulata Spiraea japonica Syzygium cumini Tamarix pentandra Triadica sebifera Ulex europaeus Waterhousea floribunda Ziziphus mauritiana

Tree/Shrub Tree/Shrub Tree/Shrub Tree/Shrub Tree/Shrub Tree/Shrub Tree/Shrub Tree/Shrub Tree/Shrub Tree/Shrub Tree/Shrub Tree/Shrub Tree/Shrub Tree/Shrub Tree/Shrub Tree/Shrub Tree/Shrub Tree/Shrub Tree/Shrub Tree/Shrub Tree/Shrub Tree/Shrub Tree/Shrub Tree/Shrub Tree/Shrub Tree/Shrub Tree/Shrub Tree/Shrub Tree/Shrub Tree/Shrub

www.issg.org/database/species/ecology.asp?si=41&fr=1&sts= www.issg.org/database/species/ecology.asp?si=100&fr=1&sts= van Wilgen et al., 2001 www.issg.org/database/species/ecology.asp?si=440&fr=1&sts= van Wilgen et al., 2001 www.issg.org/database/species/ecology.asp?si=43&fr=1&sts= www.issg.org/database/species/ecology.asp?si=332&fr=1&sts= www.issg.org/database/species/ecology.asp?si=344&fr=1&sts= www.issg.org/database/species/ecology.asp?si=261&fr=1&sts= Starfinger et al., 2003 www.issg.org/database/species/ecology.asp?si=59&fr=1&sts= www.issg.org/database/species/ecology.asp?si=211&fr=1&sts= www.issg.org/database/species/ecology.asp?si=809&fr=1&sts= www.issg.org/database/species/ecology.asp?si=212&fr=1&sts= www.issg.org/database/species/ecology.asp?si=572&fr=1&sts= www.issg.org/database/species/ecology.asp?si=215&fr=1&sts= www.issg.org/database/species/ecology.asp?si=79&fr=1&sts= www.issg.org/database/species/ecology.asp?si=476&fr=1&sts= www.issg.org/database/species/ecology.asp?si=500&fr=1&sts= www.issg.org/database/species/ecology.asp?si=22&fr=1&sts= van Wilgen et al., 2001 www.issg.org/database/species/ecology.asp?si=209&fr=1&sts= www.issg.org/database/species/ecology.asp?si=75&fr=1&sts= www.issg.org/database/species/ecology.asp?si=622&fr=1&sts= www.issg.org/database/species/ecology.asp?si=505&fr=1&sts= Pimentel et al., 2000; www.issg.org/database/species/ecology.asp?si=72&fr=1&sts=sss www.issg.org/database/species/ecology.asp?si=712&fr=1&sts= www.issg.org/database/species/ecology.asp?si=69&fr=1&sts= www.issg.org/database/species/ecology.asp?si=483&fr=1&sts= www.issg.org/database/species/ecology.asp?si=214&fr=1&sts=

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Annex 2. Select alien invasive species affecting forests and forestry


The Invasive Species Specialist Group (ISSG) of the Species Survival Commission (SSC) of the World Conservation Union (IUCN) has compiled a list of the One Hundred of the Worlds Worst Invasive Alien Species (www.issg.org/database/species/search.asp?st=100ss&fr=1&sts=#SpeciesList) which aims to collectively illustrate the range of impacts caused by biological invasion. Although incomplete, this list is a first attempt to rank the impact of alien invasive species. Included in this list are 62 alien invasive species that impact forests and forestry. This annex provides an overview of these species. Unless otherwise noted, the information in this annex is taken from the Global Invasive Species Database developed by ISSG (www.issg.org/database [accessed on 23 August 2005]). More detailed information on these species can be found there.

Fungi
Batrachochytrium dendrobatidis is a parasitic fungus that causes fungal skin infections (chytridiomycosis) in wild and captive amphibians which has been associated with population declines in endemic amphibian species in montane rain forests (Daszak et al., 1999; Weldon et al., 2004). Chytridiomycosis has been reported in 38 amphibian species from 12 families, including frogs, toads and salamanders.
Although the fungus has been found on every continent that has amphibians except Asia, Africa is suggested as the origin of B. dendrobatidis (Speare and Berger, 2000; Weldon et al., 2004). The fungus can be found in lakes, natural forests, riparian areas and water courses and can remain viable in the environment, especially aquatic environments, for weeks.

Cryphonectria parasitica, known commonly as chestnut blight, is a fungus that primarily attacks chestnut

trees (Castanea spp.) but can also cause damage to oak trees (Quercus spp.) and other hardwood tree species. It occurs in natural and planted forests where it infects the above-ground parts of trees only, creating cankers that expand, girdle and eventually kill tree branches and trunks. The native range of C. parasitica is Asia and it is considered invasive in Europe and North America.

First observed killing trees in the Bronx Zoo in New York City in 1904, the fungus was likely introduced to North America on nursery stock from Asia (Canadian Chestnut Council, undated). From there, chestnut blight spread throughout eastern North America and by 1940, three and half billion chestnut trees had perished (Cock, 2003). Dutch elm disease is a wilt disease caused by a pathogenic fungus, Ophiostoma ulmi sensu lato, which is spread by specialized bark beetles that breed under the bark of dying elm trees (Stack, McBride and Lamey, 1996). The young adult beetles fly from infected pupal chambers to feed on healthy elm trees carrying fungal spores on their bodies and depositing them in healthy plant tissue. The fungus can also spread via root grafts. The symptoms and progression of the disease differ among trees which are infected through beetle feeding and those which are infected through root grafts (Stack, McBride and Lamey, 1996). Trees infected by beetles first show wilting, curling and yellowing of leaves on one or more branches in the upper crown of the tree. Large trees may survive and progressively show more symptoms for years after initial infection. Trees infected through root grafts tend to wilt and die rapidly as the disease progresses from the base of the tree upward, typically in the spring after leaves have developed. 40

During the last century, there have been two destructive pandemics of this disease in Europe and North America, caused by the successive introduction of two fungal pathogens: Ophiostoma ulmi and the much more aggressive, Ophiostoma novo-ulmi. The fungus occurs in natural and planted forests and urban areas. Host trees include all the Euro-American native elms such as field elm (Ulmus minor), wych elm (Ulmus glabra), European white elm (Ulmus laevis), American elm (Ulmus americana), red or slippery elm (Ulmus rubra), rock elm (Ulmus thomasii) and cedar elm (Ulmus crassifolia). Asian elm species are generally much less susceptible than EuroAmerican native elms and although the Chinese elm (Ulmus parvifloia) is occasionally infected by the disease, it is much less susceptible than American elm.

Phytophthora ramorum is the causal agent of sudden oak death which until 2000 was undiscovered and unnamed. This fungus is the cause of much concern in Europe and North America because of the high level of local destruction it causes, the lack of knowledge of its epidemiology, and its high prevalence in nurseries which increases the potential of spread to new areas (Garbelotto, 2004).
P. ramorum occurs in natural and planted forests and has an extensive host range, covering many plant genera and several families, including trees, shrubs and woody and herbaceous perennials. The hosts can be broken down into two categories - bark canker hosts and foliar hosts - depending on the part of the plant or tree most susceptible to the disease (Rizzo and Garbelotto, 2003; Garbelotto, 2004). Bark canker hosts become infected on their trunks and in highly susceptible species, the cankers eventually girdle the tree, resulting in starvation and death. Mortality may occur in as little as two years. Foliar hosts become infected on their leaves and twigs and may be accompanied by branch infections and dieback in some foliar hosts; such hosts only occasionally die from infection. The fungus spreads to new locations primarily through the nursery trade and are locally spread by vectors, and infected soil, water and tools. Infestations of P. ramorum have been reported in nurseries in most European countries, the United States (Oregon and Washington) and Canada (British Columbia) (Garbelotto, 2004). It is unknown how this fungus originally entered Europe and North America, but genetic analysis suggests the pathogen is from a third location (Rizzo et al., 2002; Garbelotto, 2004).

Flatworm
introduced to many islands throughout the Pacific and Indian Oceans as a biological control agent for the giant African snail (Achatina fulica) (Lydeard et al., 2004). It now poses a serious threat to native gastropods, including many endemic snail species. This flatworm occurs in agricultural areas, coastland, disturbed areas, natural forest, planted forests, riparian zones, scrublands, shrublands, urban areas, and wetlands though is generally found in leaf litter in both undisturbed forests and habitats heavily modified by humans. Originating from New Guinea, P. manokwari now occurs in Australia, Guam, Hawaii, Japan, Maldives, Northern Mariana Islands, Palau, Philippines, Samoa, Tonga and Vanuatu (Lydeard et al., 2004).

Platydemus manokwari is a predatory species of flatworm that has been intentionally and unintentionally

Insects
Aedes albopictus, the Asian tiger mosquito, was introduced to the United States and other countries via
used tire imports. This mosquito attacks more hosts than any other mosquito in the world, including many mammals, birds and reptiles, and is associated with the transmission of dengue fever, yellow fever, eastern equine encephalitis, dog heartworm and possibly St. Louis and LaCrosse encephalitis viruses (Wittenberg and Cock, 2001). A. albopictus occurs in natural forests and urban areas.

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Believed to be indigenous to West Africa, Anoplolepis gracilipes or the yellow crazy ant, has been introduced, primarily through cargo shipments, into Africa, Asia, Australia, South America and many islands in the Caribbean, Indian and Pacific Oceans (Holway et al., 2002; Matthews, 2004). In some cases, however, the ant was intentionally introduced as a biological control agent against insect pests in cocoa, coconut and coffee plantations (Matthews, 2004). Because of its mutualistic relationship with Homoptera, A. gracilipes has itself become a pest of the agriculture and forest sectors. In exchange for their sugary honeydew secretions, the ant protects scale insects from predators and spreads them to new host plants. At high densities, these scale insects weaken plants and, combined with a sooty mould that inhabits the honeydew, can cause canopy dieback or even the death of the tree (Secretariat of the Convention on Biological Diversity, 2003b; Matthews, 2004). The yellow crazy ant affects native fauna by competing for food resources, altering the habitat or direct predation. In at least one case its removal of a keystone species, the red land crab on Christmas Island, has resulted in a significant change in forest composition and structure and a decrease in nutrient cycling (Secretariat of the Convention on Biological Diversity, 2003b; Matthews, 2004). A. gracilipes can also affect the reproductive success of many mammals, reptiles and birds, such as the Christmas Island thrush (Turdus poliocephalus erythropleurus) (Secretariat of the Convention on Biological Diversity, 2003b). Such impacts have also affected the tourism sector in many areas. For example, on Bird Island in the Seychelles, the ant has displaced approximately 60 000 pairs of sooty terns (Sterna fuscata), which are a major tourist attraction of the island (Secretariat of the Convention on Biological Diversity, 2003b). A. gracilipes occurs in agricultural areas, coastland, disturbed areas, natural and planted forests, rangelands, grasslands, riparian zones, scrublands and shrublands, urban areas and water courses. The Asian longhorned beetle, Anoplophora glabripennis, native to China and Korea, occurs in agricultural areas, disturbed areas, natural and planted forests, scrublands and shrublands, and urban areas. They attack hardwood trees, feed on the leaves, petioles and twigs and then bore into the tree trunk to lay their eggs (Matthews, 2004). After hatching, the larvae tunnel under the bark and feed on plant tissue until they are ready to pupate. Adult beetles emerge through small exit holes. Repeated attacks cause tree dieback and eventually kill the tree, typically over an estimated 3-5 year period, though some tree species may live longer (Matthews, 2004). Infestations were first discovered in North America in New York City in 1996 and in Chicago in 1998, about ten years after the beetle first entered the United States. As of May 2001, 5 286 and 1 547 infested trees have been cut in New York and Chicago respectively. An infestation was also discovered in Austria in 2001, an estimated 2-3 years after its introduction, and as of August 2001, approximately 47 infested trees have been felled. Asian longhorned beetles were also noted in New Jersey, United States in 2002 and in an industrial park in the Toronto-Vaughan area, Canada in 2003. In Poland, a single specimen was found on a bonsai tree (Acer palmatum) in 2003 and the beetles were also discovered in France in 2003 and 2004, about 100 km south of Paris and near Nante Port. In its native China, Asian longhorned beetles have damaged about 40 percent (approximately 2.3 million ha) of poplar plantations - over 240 cities or counties have been infested in five provinces alone (totalling 230 thousand ha) - and an estimated 50 million trees were cut down over a three year period (1991-1993) in Ningxia Province alone.

Cinara cupressivora, the cypress aphid previously referred to as C. cupressi, is a brownish soft-bodied insect that has been observed around the world feeding on various trees species from several genera including Austrocedrus, Callitris, Chamaecyparis, Cupressus, Juniperus, Thuja, Widdringtonia and the hybrid Cupressocyparis (Watson et al., 1999; Mwangi, 2002). The species occurs in natural and planted forests where it sucks the sap

42

from twigs causing yellowing to browning of foliage. Depending on the severity and duration of the infestation, the overall effects range from partial damage to eventual tree death (O'Neil, 1998). The cypress aphid has critically damaged commercial and ornamental plantings and native stands of Cupressus, Juniperus, Widdringtonia and other Cupressaceae in Africa, Colombia, Italy, Jordan, Mauritius and Yemen (Watson et al., 1999). Up to 1990, it was estimated that C. cupressivora had killed approximately US$44 million worth of trees throughout southern and eastern Africa and was causing an additional loss of approximately US$14.6 million per year through reductions in annual growth increment (Cock, 2003). In Kenya, approximately 5 153 ha of 86 000 ha of industrial forest plantations of Cupressus lusitanica have been infested by the aphid to varying degrees ranging from slight to severe (Mwangi, 2002). A study conducted by the Kenya Forestry Institute estimated that the cypress aphid in Kenya might kill as many as 50 percent of all cypress trees during the 30-year harvest cycle (Cock, 2003). C. cupressivora probably originated in a region from eastern Greece to just south of the Caspian Sea and is now known to exist in Africa, Asia, Europe, North America and South America (Watson et al., 1999).

Linepithema humile, commonly known as the Argentine ant, is a very successful colonizer that produces

large numbers of aggressive workers capable of invading both disturbed and undisturbed habitats. In favourable environments, the Argentine ant can attain high densities and displace native and introduced ant species, significantly reduce the abundance of other arthropods, and alter ecosystem level processes such as plant pollination and seed dispersal (Cole et al., 1992; Human and Gordon, 1996; Human and Gordon, 1997; Holway, 1998; Christian, 2001). The native range of the Argentine ant is South America and the species can now be found on all continents except Antarctica, as well as a number of oceanic islands (Suarez, Holway and Case, 2001). Southern Europe is home to the world's largest colony of L. humile. It occurs in agricultural areas, coastal areas, disturbed areas, natural and planted forests, rangelands and grasslands, riparian zones, scrublands and shrublands, urban areas and wetlands.

The gypsy moth, Lymantria dispar, is one of the most destructive pests of hardwood forests and shade, fruit and ornamental trees throughout the northern hemisphere. The caterpillars cause extensive defoliation, leading to reduced growth or even death of the host tree. They destroy the aesthetic beauty of natural areas by defoliating and killing the trees and covering the area with their waste products and silk. Gypsy moth also has impacts on human health as the hairs on larvae and egg masses have been known to cause allergies in some people. Cyclical outbreaks generally only occur in stands where primary hosts, such as Quercus and Populus, comprise more than 20 percent of the basal area and where environmental conditions are favourable to generation explosions. Outbreaks (of several generations) can be every 7-10 years, however more recently the period between outbreaks is apparently becoming shorter. L. dispar can be found in natural and planted forests, riparian zones and urban areas. The native range of the European gypsy moth is southern Europe and northern Africa. It was introduced to the United States by a scientist searching for silk-producing caterpillars (Wittenberg and Cock, 2001; Ciesla, 2002). This form soon moved to Canada and is now present in most of northeastern North America and its range is expanding to the south and west. The native range of the Asian gypsy moth is central and southern Asia and Japan. This form is considered a greater threat as adult females are capable of flight and can therefore disperse greater distances. It is classified as a potential threat by many countries and is covered by phytosanitary regulations.

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Regular monitoring, generally through the use of pheromone traps, is carried out to intercept establishment of the gypsy moth. having attained a pantropical distribution. It is a serious threat to biodiversity as it aggressively displaces most native invertebrate species (Hoffmann, Andersen and Hill, 1999). Reductions in vertebrate populations have also been observed where this ant is extremely abundant. The big-headed ant is also a domestic pest since it chews on telephone cabling and electrical wires and an agricultural pest as it harbours phytophagous insects that reduce crop productivity (Hoffmann, Andersen and Hill, 1999). P. megacephala can also facilitate the invasion of non-native plant species. P. megacephala occurs in agricultural areas, coastal areas, disturbed areas, natural and planted forests, rangelands, grasslands, riparian zones, scrublands and shrublands, urban areas and wetlands. The native range is believed to be southern Africa and it is now widespread throughout the temperate and tropical zones of the world. It favours shaded and moist environments but can exist wherever there is anthropogenic disturbance (Hoffmann, Andersen and Hill, 1999). Red imported fire ants (RIFA), Solenopsis invicta, are aggressive generalist foragers that occur in very high densities thereby allowing them to dominate food sources. Their stinging ability allows them to subdue prey and repel even large vertebrate competitors from resources. They have considerable impacts on native invertebrates. In the southeastern United States, for example, S. invicta has reduced native ant abundance by 90 percent and species richness by 70 percent and has had a serious impact on other invertebrates (Hoffmann, Andersen and Hill, 1999). This species breeds and spreads rapidly and if disturbed, can relocate rapidly in order to ensure colony survival. Native to South America, the red imported fire ant has been introduced into parts of Australia, New Zealand, North America and some Caribbean islands, notably Puerto Rico and the Virgin Islands (Holway et al. 2002). S. invicta occurs in agricultural areas, coastal areas, deserts, disturbed areas, natural and planted forests, rangelands, grasslands, riparian zones, scrublands, shrublands, urban areas and water courses. S. invicta inhabits hot arid regions however it may survive colder climates in human habitations or infrastructure such as climate-controlled buildings or greenhouses. Invasive ants are usually more likely to establish in disturbed habitats, including the edges of forests or agricultural areas (Ness and Bronstein, 2004). Deforested areas are particularly at risk of being invaded by red imported fire ants (Morrison et al., 2004). This ant constructs earthen mounds for brood thermoregulation, which are easier to build in open, sunny areas; as a result it is less abundant in, and poses a smaller threat to, dense forests (Morrison et al., 2004). Tropical regions that are warm, wet and densely forested are not suitable habitats for fire ants (Morrison et al., 2004).

Pheidole megacephala, commonly called the big-headed ant, is one of the most invasive species of ant,

Vespula vulgaris, the common wasp or yellowjacket, occurs in natural and planted forests, scrublands and

shrublands and urban areas where they nest underground and in tree cavities and buildings. The common wasp is a Holarctic species and has been introduced into New Zealand and Australia. In addition to causing painful stings to humans, they compete with birds and other insects for insect prey and sugar sources. They will also eat fruit crops and scavenge around rubbish bins and picnic sites. This species impacts a wide range of sectors including conservation, forestry, beekeeping, horticulture, and human health. V. vulgaris is a major threat to insect and bird populations in native beech forests on New Zealands South Island (Matthews et al., 2000). introduced into parts of Africa, North America, South America and onto some islands in the Caribbean and the Pacific Oceans including the Galapagos, Hawaii, New Caledonia, Vanuatu and the Solomon Islands (Roque-Albelo and Causton, 1999; Holway et al., 2002). The ant occurs in agricultural areas, coastlands,

Wasmannia auropunctata, or the little fire ant, is native to Central and South America and has been

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deserts, disturbed areas, natural and planted forests, rangelands and grasslands, riparian zones, scrublands and shrublands, urban areas and wetlands. The little fire ant reduces species diversity and abundance of flying and tree-dwelling insects and eliminates arachnid populations. In agricultural areas, the ant is considered a great nuisance to humans because it is more likely to reach high densities and sting people working in the field. In human habitations, it may sting and even blind domestic pets (Romanski, 2001). In New Caledonia and the Galapagos, W. auropunctata has been blamed for a decrease in reptile populations, where it eats tortoise hatchlings and attacks the eyes and cloacae of adult tortoises (Nishida and Evenhuis, 2000). The ant is probably the most aggressive species that has been introduced into the Galapagos where a marked reduction of scorpions, spiders and native ant species in infested areas has been observed (Roque-Albelo and Causton, 1999; Nishida and Evenhuis, 2000). The species has also led to a decrease in local arthropod biodiversity in the Solomon Islands (Romanski, 2001). In its native range the little fire ant decreases herbivorous arthropods, increasing plant growth and fruit and seed production, and decreasing pathogen attacks. W. auropunctata may also however exclude arthropod plant mutualists such as plant tenders or seed dispersers (Ness and Bronstein, 2004). It is considered to be perhaps the greatest ant species threat in the Pacific Region. Environmental stresses, including human practices such as monoculture plantings, may cause explosions of some ant populations. For example, in its native range in South America, W. auropunctata is a pest in disturbed forests and agricultural areas where it can reach higher densities. High densities of this species have been linked with sugar cane monocultures in Colombia and cocoa farms in Brazil. In Colombia, a high abundance of the little fire ant in forest fragments has also been linked with low native ant diversity. The little fire ant efficiently exploits resources including shelter, nectar, and honeydew residues of Homopteran insects and may outcompete and displace native ants (Armbrecht and Ulloa-Chacn, 2003).

Molluscs
Achatina fulica, the giant African snail, is native to East Africa and has been introduced into most regions of the humid tropics, including many Pacific islands, eastern and southern Asia, and the Caribbean. It has also been intercepted widely by quarantine officials and initial invasions have been eradicated in many countries. The species occurs in agricultural areas, coastal areas and wetlands, disturbed areas, natural and planted forests, riparian zones, scrublands and shrublands, and urban areas
A. fulica has been considered the most important snail pest in the tropics and subtropics and perhaps the world. A major agricultural and garden pest, and a general nuisance, the giant African snail eats native plants, modifies habitats and out-competes native snails (Matthews, 2004). A. fulica is also a vector for eosinophilic meningitis, a disease caused by the parasite rat lungworm that is passed to humans through the consumption of raw or improperly cooked snails (Matthews, 2004). Often the introduction of the giant African snail has lead to the subsequent introduction of predatory snails and flatworms as biological control agents that can also have devastating effects on native land snail diversity. The predatory rosy wolf snail, Euglandina rosea, is native to the southeastern United States and has been introduced to islands in the Caribbean, Pacific and Indian Oceans as a biological control agent for another alien invasive species, the giant African snail (Achatina fulica) (Stein and Flack, 1996). This snail has caused the extinction of numerous endemic tree snails in French Polynesia and has been heavily implicated in the extinction or decline of other snail species wherever it has been introduced, notably in Hawaii (Stein and Flack, 1996). E. rosea occurs in disturbed areas, natural and planted forests, scrublands and shrublands, and urban areas.

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Amphibians
Bufo marinus, or cane toads, are indigenous to northern South America, Central America, and Mexico northward to the southern United States and have been introduced throughout the world as a biological control agent for various insect pests of sugarcane and other crops. In its introduced range, it has become a serious pest feeding on a wide variety of insects, frogs, reptiles, birds and mammals (Department of Natural Resources and Mines, 2003). It also preys on and competes with native amphibians for food and breeding habitat. Glandular secretions excreted when the toad is provoked or pressure is applied, are toxic and are known to cause illness and death in domestic animals and wildlife that come into contact with toads. B. marinus inhabits agricultural areas, disturbed areas, lakes, water courses, wetlands and riparian zones, natural forests and urban areas. Eleutherodactylus coqui is a small tree frog native to Puerto Rico that has been introduced into the AsiaPacific region, North America and the Galapagos Islands. The Caribbean tree frog is highly adaptable to different ecological zones and altitudes and inhabits agricultural areas, natural and planted forests, wetlands and riparian zones, and urban areas.

Their loud call is the main reason they are considered pests and in some regions there are fears that this may negatively impact the tourism industry (HEAR, 2004). These frogs have a voracious appetite and in Hawaii, there is concern that E. coqui may put endemic insect and spider species at risk and compete with endemic birds and other native fauna which rely on insects for food (Campbell, 2000).

Reptiles
Boiga irregularis, the brown treesnake, is native to eastern Indonesia, the Solomon Islands, New Guinea, and the northern and eastern coasts of Australia and has been introduced to Guam and the Northern Mariana Islands. They have also been sighted on Saipan, Tinian, Rota, Kwajalein, Wake Oahu, Pohnpei, Okinawa, and Diego Garcia. The brown treesnake occurs in agricultural areas, wetlands, riparian and coastal zones, disturbed areas, natural and planted forests, rangelands and grasslands, scrublands and shrublands, and urban areas.
The brown snake is notorious for concealing itself in international freight and aircraft. It is nocturnal, secretive and arboreal, hunting for food at all levels within a forest. It is an effective generalized predator that has decimated the avifauna, greatly impacted other terrestrial vertebrates, caused cascading ecological perturbations, and disrupted the electrical supply in Guam.

Birds
Pycnonotus cafer is a noisy, gregarious bird that occurs naturally from Pakistan to southwest China and has
been introduced to many Pacific Islands where it is considered highly invasive. Commonly known as the redvented bulbul, it is an agricultural pest that destroys fruits, flowers, beans, tomatoes and peas. It may also help to spread the seeds of other alien invasive species. The bulbul is aggressive and chases off other bird species. It occurs in agricultural areas, natural and planted forests, scrublands and shrublands, and urban areas.

Sturnus vulgaris, the European starling, is an aggressive omnivore native to Asia, Europe and North Africa
and has been introduced into Australia, New Zealand, North America and South Africa. The starling prefers lowland habitats to more mountainous terrain and inhabits agricultural areas, coastal areas, disturbed areas, natural and planted forests, rangelands, grasslands, riparian zones and wetlands, scrublands and shrublands, tundra and urban areas.

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European starlings cost hundreds of millions of dollars in agricultural damage each year and contribute to the decline of local native bird species through competition for resources and nesting space. They are also a public nuisance, can damage infrastructure, negatively affect aesthetics, and can pose public heath risks by transmitting diseases to humans as well as livestock (Adeney, 2001).

Mammals
Goats, Capra hircus, are destructive herbivores with a highly diverse diet that includes plant species often avoided by sheep or cattle. They prevent successful regeneration of favoured plant species and their overbrowsing also decreases biomass and percent cover of vegetation, resulting in a decrease or extinction of animal species, and often severe erosion on the devegetated landscape. In New Zealand, for example, 35 years of overgrazing, overbrowsing and trampling by goats reduced the number of plant species from 143 to only 70 (Courchamp, Chapuis and Pascal, 2003). Goats easily become feral and can spread disease to native wildlife. Grasslands, scrublands and forests are all used extensively as feeding areas.

Cervus elaphus, or red deer, are a Eurasian species that have been introduced to Australia, New Zealand,

North America and South America. They inhabit natural and planted forests, rangelands and grasslands, scrublands and shrublands, and tundra. Deer prevent regeneration of preferred plant species, thereby creating significant changes to the structure and composition of native ecosystems (Department of Conservation, 2002). In some areas, non-replacement of canopy species can lead to canopy collapse. Even at low densities, red deer inhibit forest regeneration (Department of Conservation, 2002). In Argentina, they have invaded several national parks where they negatively affect native flora and fauna and possibly disrupt ecological processes (Jaksic et al., 2002). In Chile, there is a concern over red deer competition with endangered endemic deer species such as the huemul, Hippocamelus bisulcus, and the pudu, Pudu pudu (Jaksic et al., 2002). Red deer also compete with livestock.

Cats, Felis catus, occur in almost all geographical regions but are usually absent from true desert, high mountain and frozen areas due to an absence of suitable prey and wetlands and rainforests as they prefer to stay dry. Their preferred prey is small mammals, amphibians, lizards and snakes but they also prey on birds, particularly those that nest or feed on or near to the ground. They have significant impacts on the abundance of such species, even to the point of extinction, thus causing considerable ecological impacts (Courchamp, Chapuis and Pascal, 2003).

Herpestes javanicus, or mongooses, occur in agricultural areas, coastal areas and wetlands, deserts,

disturbed areas, natural and planted forests, rangelands and grasslands, riparian zones, scrublands and shrublands, and urban areas. They were introduced to many islands in the West Indies beginning in 1872 and to Fiji and several of the larger Hawaiian Islands in 1883 to control rats in sugar cane fields (Atkinson and Atkinson, 2000). Mongooses prey on birds, small mammals and reptiles and have caused the decline or extinction of many endemic vertebrates (Courchamp, Chapuis and Pascal, 2003). They cause significant losses to industries involving game species and livestock and are also a vector of rabies.

Macaca fascicularis, commonly known as the crab-eating or long-tailed macaque, is native to mainland Southeast Asia, the Philippines, and parts of Indonesia and has been introduced into Mauritius, Palau, Hong Kong and other parts of Indonesia (Tinjil Island and Papua) where it is considered to be invasive or potentially invasive. They inhabit a wide range of habitats including riparian, secondary and primary forests, coastal forests, forest edges, swamps, and urban and agricultural settings. Highly adaptive to new environments, macaques often prefer secondary habitats which have been disturbed by human activity. It is an opportunistic mammal and reaches higher densities in degraded forest areas. In its introduced ranges, macaques have few natural predators and negatively affect native biodiversity by consuming native plants and competing with birds for fruit and seed resources. They also facilitate the spread of alien invasive plants

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through seed dispersal. Macaques may impact the agriculture and forest sectors by feeding on commercially important crops, plants and trees. They may also carry potentially fatal human diseases including B-virus. The house mouse, Mus musculus, probably has the most extensive global distribution than any other mammal with the exception of humans. Native to the Indian subcontinent, the geographic spread of the house mouse has been facilitated by its commensal relationship with humans and they have colonized tropical, temperate, semi-desert, desert and subantarctic regions throughout the world. They are common in agricultural areas, coastal areas, disturbed areas, natural and planted forests, rangelands and grasslands, riparian zones, scrublands and shrublands, and urban areas. House mice are major economic pests; consuming, destroying and contaminating crops and food supplies intended for human consumption. They have also been implicated in the extirpation or extinction of many indigenous species. They are host to a range of human diseases and parasites, the most serious being bubonic plague (Yersinia pestis) and salmonella (Salmonella spp.). introduced into New Zealand and some small European islands. They live anywhere they can find suitable prey but prefer forests or shrublands to protect them from their avian predators. They also inhabit agricultural areas, coastal areas, disturbed areas, riparian zones, scrublands and shrublands, tundra and wetlands. An intelligent, resourceful predator, the stoat specializes in small mammals and birds though they also attack animals larger than themselves. They have contributed to the loss of native birds in New Zealand, including the northern brown kiwi, Apteryx mantelli, and hole-nesting forest birds (O'Donnell, 1996; Basse, McLennan and Wake, 1999). Research and management of stoats in New Zealand costs millions of dollars every year. Native to southern Europe and North Africa, rabbits, Oryctolagus cuniculus, have been introduced to all continents except Antarctica and Asia. They are found in agricultural areas, desert, disturbed areas, natural and planted forests, rangelands and grasslands, scrublands and shrublands, and urban areas. Rabbits cause considerable damage to the environment and agricultural areas. They compete with native wildlife for food and shelter, contribute to a decline of many native plants and animals, and by supporting large populations of predators such as cats and foxes, they can also enhance their negative impacts on native species (Courchamp, Chapuis and Pascal, 2003). Rabbits cause extensive erosion through browsing and loss of plant cover and often destroy the habitats of many animals (Courchamp, Chapuis and Pascal, 2003). They also compete with livestock for food.

Mustela erminea, or stoat, is native to northern parts of Eurasia and North America and has been

Rattus rattus is a native of the Indian subcontinent that has spread worldwide inhabiting agricultural areas,

coastal areas, disturbed areas, natural and planted forests, rangelands and grasslands, riparian zones, scrublands and shrublands, and urban areas. Ship rats are omnivorous and capable of eating a wide range of plants and animals including the eggs and young of forest birds (Innes, 2001; Courchamp, Chapuis and Pascal, 2003; Robertson and Saul, 2004). They are known to impact the regeneration of maritime forest due to their consumption of inland podocarp seeds and their competition with and predation of many bird species that disperse seeds (Courchamp, Chapuis and Pascal, 2003). They have directly caused or contributed to the extinction of many species of wildlife including birds, small mammals, reptiles, invertebrates and plants, particularly on islands. The ship rat is the species most frequently identified with catastrophic declines of birds on islands as evidenced on Midway Island in the Leeward Islands of Hawaii, Lord Howe Island, and Big South Cape Island, New Zealand (Atkinson and Atkinson, 2000; Courchamp, Chapuis and Pascal, 2003). They are known to prey on many bird species including the endangered kakerori (Pomarea dimidiata) in the Cook Islands, sooty terns in the Seychelles, Bonin petrels in Hawaii, Galapagos dark-rumped petrels in the Galapagos Islands, and white-tailed tropicbirds in Bermuda (Atkinson and Atkinson, 2000; Robertson and Saul, 2004).

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Native to the eastern United States, grey squirrels, Sciurus carolinensis, were imported into Italy, South Africa and the United Kingdom for the pet industry. They have subsequently been released or have escaped into the wild and are known to live in agricultural areas, disturbed areas, natural and planted forests, scrublands and shrublands, and urban areas. In the United Kingdom and Italy, the grey squirrel is expanding its range resulting in a decrease of the native red squirrel. It causes considerable damage to woodlands by stripping the bark from trees. The grey squirrel is also suspected to be a vector for the parapoxvirus which is fatal to red squirrels. Feral pigs, Sus scrofa, are escaped or released domestic animals that are now found in many parts of the world. They cause significant damage to crops and property and transmit many diseases to domestic and wild animals. Pigs damage forests by eating or uprooting tree and plant seedlings and by opening tree-fern trunks in searching for starch (Atkinson and Atkinson, 2000). Pigs can spread alien invasive weeds by feeding on some species of fruit such as guava; the seeds pass through the gut and into droppings (Atkinson and Atkinson, 2000). They can also affect large native invertebrates, such as earthworms and land snails, through direct predation and habitat destruction (Atkinson and Atkinson, 2000). Pigs are very effective predators of seabirds such as albatrosses, shags and boobies (Atkinson and Atkinson, 2000). They are suspected to facilitate infestation of trees by the dieback disease caused by the fungus, Phytophthora cinnamomi (Atkinson and Atkinson, 2000). The brushtail possum, Trichosurus vulpecula, is a solitary, nocturnal, arboreal marsupial from Australia that was introduced into New Zealand where it is causing considerable damage to native forests by selective feeding on foliage and fruits. Persistent possum browsing pressure can result in long-term changes in forest structure and composition (Courchamp, Chapuis and Pascal, 2003). This species also competes for tree hollows with endemic hole-nesting species, such as kiwi (Apteryx spp.), preys on invertebrates and the eggs and chicks of birds, and transmits bovine tuberculosis to cattle and deer (Courchamp, Chapuis and Pascal, 2003). The red fox,Vulpes vulpes, is one of the most widely distributed carnivores in the world inhabiting agricultural areas, deserts, disturbed areas, natural and planted forests, rangelands and grasslands, riparian zones, scrublands and shrublands, tundra and urban areas. Native to Asia, Europe, North Africa and the boreal regions of North America, this species has been introduced into Australia and temperate regions of North America. Red foxes are generalist predators that attain relatively high densities and negatively impact many native species, including smaller canids and ground nesting birds in North America, and many small and medium-sized rodent and marsupial species in Australia (Kamler and Ballard, 2002). They also prey on livestock. In Canada, introduced red foxes may have already replaced several native subspecies of red foxes. They also pose a threat to the health of humans and domestic animals through the transmission of diseases such as rabies, distemper, parvo virus and mange.

Grass
subtropical and temperate areas around the world. Considered native to the Indian subcontinent, the giant reed is now present in the Federated States of Micronesia, Fiji, Guam, Hawaii, Nauru, New Caledonia, Norfolk Island, Palau, Samoa and Christmas Island.

Arundo donax is a tall perennial grass that has been widely introduced into riparian zones and wetlands in

Once the reed becomes established, it forms impenetrable, homogenous stands which displaces native plant species and thus alters the habitat of native wildlife (Benton, Bell and Swearingen, undated). It is also both a fire and flood hazard. Giant reed traps sediments and narrows flood channels, leading to erosion and flooding, and promotes wildfire. It may also reduce water availability through high evapotranspiration rates.

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Native to Southeast Asia, Imperata cylindrica, or cogongrass, can flourish in virtually all ecosystem types including degraded forests, grasslands, arable land and young plantations, in both temperate and tropical regions. It is a considerable nuisance species that displaces native plant species, disrupts the habits of wildlife, and alters natural fire cycles by causing more frequent and intense fires (van Loan, Meeker and Minno, 2002). The costs of controlling cogongrass amount to hundreds of millions of dollars annually.

Plants
Commonly called chromolaena, triffid weed or Siam weed, Chromolaena odorata is a fast-growing perennial shrub native to Central and South America that has been introduced into much of Africa, tropical Asia and the Pacific. An aggressive invasive weed, it forms dense stands which prevent the establishment of other species through competition and allelopathic effects. Chromolaena can be found in agricultural areas, disturbed areas, natural and planted forests, rangelands and grasslands, scrublands and shrublands, and riparian zones. In South Africa, it is considered a considerable threat to conservation and ecotourism, as it has primarily invaded natural areas, reducing the biodiversity of grasslands, savannahs and forests, and compromising game-viewing in nature reserves and national parks (Matthews and Brand, 2004). It has also been shown to threaten the breeding habitat of the Nile crocodile in South Africa and the decrease in temperature caused by the shading and crowding of nesting sites can induce female-biased sex ratios or may even prevent embryonic development altogether (Leslie and Spotila, 2001; Matthews and Brand, 2004). In Asia, chromolaena decreases agricultural productivity by readily invading subsistence food gardens, cultivated crops, and young or neglected plantations of cocoa, coconut, oil palm, rubber and tobacco (Matthews, 2004). This weed is a considerable problem in commercial tree plantations as it suppresses the growth of young pine and eucalypt trees and allows fire to penetrate deeper into plantations (Matthews, 2004; Matthews and Brand, 2004). It can also promote wildland fires (PIER, 2004a; Department of Natural Resources and Mines, 2005). Chromolaena may also cause skin problems and asthma in allergy-prone people.

Euphorbia esula or leafy spurge is perennial plant native to Europe and temperate Asia that can now be found throughout the world with the exception of Australia. It occurs in agricultural areas, disturbed areas, natural forest, rangelands and grasslands, scrublands and shrublands, and urban areas. E. esula is an aggressive invader that displaces native vegetation by shading sunlight, exhausting water and nutrient supplies and excreting toxins that prevent the growth of surrounding plants. Contact with the milky sap contained in leafy spurge can have impacts on human health ranging from minor skin irritations to blindness if the liquid reaches the eyes.
Planted in many parts of the world, Hedychium gardnerianum is a native plant of India that has escaped from gardens and can now be found in the Cook Islands, the Federated States of Micronesia, French Polynesia, Hawaii, Jamaica, New Zealand, Runion and South Africa. It forms dense colonies in native forests that chokes understorey plants and prevents young native seedlings from establishing thus changing the function and structure of native forests and in some instances resulting in the death of the entire forest (Environment Bay of Plenty, 1998).

Mikania micrantha is a perennial, climbing plant that is commonly called the mile-a-minute weed because of its incredible growth rate. It grows best in areas with high fertility, organic matter, soil moisture and humidity. M. micrantha is a particular threat to young plantations and nurseries where it damages or kills other plants by blocking sunlight and suffocating them, by competing for water and nutrients, and by releasing substances that inhibit their growth (Matthews, 2004). The weed is especially problematic in tea crops in India and Indonesia and in rubber plantations in Malaysia and Sri Lanka (Matthews, 2004). It also causes serious problems in pastures and in oil palm, banana, cacao and forest plantations.

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It its native habitat in Central and South America, M. micrantha grows in and near forests, and in riparian and disturbed areas. Widely introduced as a cover crop and garden ornamental, it has since spread readily due to its efficient reproduction and can now be found in Bangladesh, India, Indonesia, Malaysia, Mauritius, Papua New Guinea, the Philippines, Sri Lanka, Thailand and many of the Pacific islands (Matthews, 2004).

Pueraria montana var. lobata, commonly known as kudzu, is a semiwoody vine with a twining and trailing growth habit that forms dense infestations that covers the ground and trees. A native of Asia, kudzu has spread and established in temperate zones and higher altitudes in the tropics and can be found growing in almost all ecosystem types including agricultural areas, disturbed areas, natural and planted forests, rangelands and grasslands, riparian areas, scrublands and shrublands, and urban areas. In the eastern United States, kudzu has infested 2 to 3 million hectares of land costing approximately US$500 million per year in land productivity losses and control costs.
Native to the tropical areas of Central America, Sphagneticola trilobata is a creeping herb that has established in many wet tropical areas worldwide. Often planted as an ornamental plant and for groundcover, it can rapidly escape from gardens to roadsides, riparian areas, natural forests, grasslands, scrublands, and plantations where it forms a dense ground cover that crowds out and prevents regeneration of other plant species (PIER, 2004b). In plantations, it competes with crops for nutrients, light and water, and reduces crop yields.

Trees and shrubs


commercial source of tannin and as a source of fuelwood and timber for local communities (Matthews and Brand, 2004). A serious threat to native habitats and species, this tree competes with indigenous vegetation, replaces grass communities, reduces native biodiversity and increases water loss from riparian zones. Native to Australia, black wattle can now be found in Africa, Asia and the Pacific, Europe, North America and South America. It occurs in disturbed areas, rangelands and grasslands, riparian areas, urban areas, and water courses.

Acacia mearnsii, commonly called black wattle, is a fast-growing, nitrogen-fixing tree often used as a

Ardisia elliptica is a fast-growing, shade-tolerant evergreen tree that forms dense stands that prevent the establishment of other plant species. A popular ornamental plant because of its fast growth and attractive fruit, it spreads from gardens and readily invades moist disturbed forests and also undisturbed sites. A. elliptica is native to the west coast of India, Indonesia, Malaysia, New Guinea and Sri Lanka and has become established in southern Florida, Hawaii, Jamaica and Okinawa. Cecropia peltata, commonly known as the trumpet tree, is native to tropical Central and South America and
has been introduced to French Polynesia, Hawaii, Malaysia and West Africa where it has subsequently become invasive. It is known to spread and establish in disturbed areas, lava flows and forest gaps.

Cinchona pubescens is a widely planted tropical forest tree from Central and South America that has

invaded a variety of forested and non-forested habitats in French Polynesia, the Galapagos Islands, Hawaii, St. Helena and Tanzania. Its fast growth, rapid spread and dense canopies result in a replacement of native vegetation in naturally treeless environments and a significant loss of species diversity of shrubs and herbs with very few species able to grow below the canopy in forests.

Clidemia hirta is a toxic weedy shrub from Central America that has been introduced to almost all tropical islands and Southeast Asia. An aggressive invader in rangelands and grasslands, scrublands and shrublands, disturbed areas and natural forests, it grows up to two metres tall forming almost impenetrable thickets which thus shade out all other vegetation (Wester and Wood, 1977).

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A native of temperate and tropical Asia, Hiptage benghalensis is a high-climbing vine to large shrub that has been cultivated in the tropics as an ornamental and for medicinal purposes (GRIN-CA, 2002; Starr, Starr and Loope, 2003). It has been a problem species in parts of Australia and the United States and is considered an extremely invasive species on the Mascarene Islands. On the islands of Mauritius and Runion, H. benghalensis flourishes in drier lowland forests and forms dense thickets that smother native vegetation and choke trees. Widely grown as an ornamental shrub throughout tropical, subtropical and temperate zones, Lantana camara has established in these regions and become a major weed of pastures, roadsides, wastelands and plantations (Matthews, 2004; Matthews and Brand, 2004). L. camara has many beneficial uses: it is grown as a hedge plant; its stalks are used for paper pulp; its bark is used as an astringent; and its leaves are used for many medicinal purposes. It also has many negative impacts on the environments in which it invades including: decreases in pasture productivity; cattle can be poisoned when lantana is ingested; exclusion of understorey species; and changes in wildlife composition through the provision of perch sites and cover (Matthews, 2004; Matthews and Brand, 2004). Access problems are also often experienced which affects recreation and forest harvesting activities. Central America (Matthews, 2004; Matthews and Brand, 2004). Subspecies leucocephala was introduced to Asia over two centuries ago and is now established around the world (Matthews, 2004). Subspecies glabrata was widely introduced across the tropics in the 1970s and 1980s by agroforestry groups and organizations and is now widely cultivated throughout the tropics and subtropics (Matthews, 2004). Leucaena is considered a conflict tree because although it is invasive and destructive, it can also provide many positive benefits. Widely promoted as a miracle tree, leucaena has been planted for the provision of wood and non-wood products and for reforestation purposes. Leucaena is spreading naturally and is considered a weed in more then 20 countries across all continents except Europe and Antarctica. It invades open, often coastal or riverine, habitats, disturbed sites and agricultural lands. It forms dense thickets which in some areas are replacing native forests and threatening endemic species of conservation concern. Such dense thickets also render extensive areas of land unusable and inaccessible, and once established the species is very difficult to eradicate. Mascarene Islands Mauritius, Runion and Rodrigues (Lavergne, Rameau and Figier, 1999; Milne and Abbott, 2004). Commonly known as Sri Lankan privet, this species exhibits rapid growth rates, high tolerance to shade and high seedling recruitment, and depends on birds to distribute its seeds. It can invade primary forests which poses a serious threat to native ecosystems (Lavergne, Rameau and Figier, 1999). Its dense foliage reduces the amount of light reaching the forest floor and prevents the regeneration of light-demanding plants. L. robustum can also alter forest structure and composition by upsetting nutrient and water cycles and by competing with native species for space and nutrients, thus displacing them and affecting successional patterns. The benefits of L. robustum are diverse which makes this a conflict species. In its native range, its stems are made into tool handles and the plant reduces erosion along streams and rivers. It was introduced into Rodrigues and Runion as a hedge plant and into Mauritius as an ornamental plant as well as to protect conifer plantations from deer, provide firewood and control invasions of other invasive weeds (Lavergne, Rameau and Figier, 1999).

Leucaena leucocephala is a fast growing, nitrogen-fixing and drought-tolerant tree native to Mexico and

Ligustrum robustum subspecies walkeri is native to Sri Lanka and is now present on all three of the

Melaleuca quinquenervia, commonly known as melaleuca or the paper bark tree, is native to Australia, New Caledonia and New Guinea. It was originally introduced into South Florida at the beginning of the twentieth century as an ornamental and to provide a useful crop capable of growing in an area subject to
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drought, flooding and periodic fires (Cock, 2003). It has since invaded across much of the state and has the potential to invade all of that region's wetlands within the next 50 years (OTA, 1993; FAO, 2000a). Melaleuca is rapidly degrading the wetlands system of the Florida Everglades by outcompeting indigenous plants and altering hydrology, topography and soils (OTA, 1993). Its dense impenetrable stands provide poor habitat and forage for wildlife and the altered hydrology has decreased populations of nesting wading birds and accelerated the proliferation and spread of alien fishes and aquatic plants (OTA, 1993). Melaleuca has also replaced sawgrass marshes, forests and other natural habitats (OTA, 1993). ornamental that, facilitated by hurricanes, feral pigs and hikers, has subsequently invaded other areas. It invades the understorey of native rain forests, eventually growing through the canopy and suppressing native trees (Denslow, 2002). In Tahiti, M. calvescens has homogenized the landscape as Miconia stands replace native forest, create barren understoreys and facilitate erosion on steep slopes (Denslow, 2002). It has spread over several of the Hawaiian Islands and the state is now spending US$1.5 million per year to protect its native rain forests (Denslow, 2002). Native to Mexico, Central and South America, Mimosa pigra has been introduced to Australia and many countries in both Africa and Asia. The giant mimosa is an aggressive woody shrub that grows rapidly, produces large quantities of seeds, and flourishes in both aquatic and terrestrial environments, allowing it to invade watercourses, floodplains and wetlands (Matthews, 2004). It forms impenetrable, prickly thickets up to four or five metres high which make invaded areas inaccessible to humans and wildlife, block irrigation channels, cause silting of dams, interfere with recreational uses of waterways, and create safety hazards along roadways. In Northern Australia, M. pigra covers about 450 km2 of river floodplain and swamp forest and these thickets have been observed to contain fewer birds and lizards, less herbaceous vegetation and fewer tree seedlings than native areas free of this species. It also prevents traditional food gathering by Aborigines and threatens pastoral industries (Binggeli, 1997). Islands and has been introduced to Australia, New Zealand and the United States, including Hawaii. A fast growing tree whose dispersal is facilitated by introduced frugivorous birds, the fire tree rapidly forms dense stands and negatively impacts the recruitment and survival of native plant species in volcanic sites, agricultural areas, disturbed areas, forests and shrublands (Walker and Vitousek, 1991). Capable of fixing nitrogen, the fire tree has been shown to alter primary successional ecosystems by increasing nitrogen levels in the soils (Vitousek, 1990). Native to the Mediterranean Basin, Pinus pinaster or maritime pine was widely planted in temperate regions within and outside its natural range for a variety of purposes. It readily regenerates almost everywhere it has been planted and has escaped and established in agricultural areas, disturbed areas, natural forests, rangelands and grasslands, riparian zones, scrublands and shrublands, and urban areas. After fire, it also regenerates profusely often resulting in dense thickets that suppress native plants, change fire regimes and hydrological properties, and alter wildlife habitats (van Wilgen et al., 2001). Native to Brazil, Psidium cattleianum or strawberry guava, has been introduced into the Mascarene Islands, Norfolk Island, Polynesia and the United States (Florida). It occurs in agricultural areas, coastal areas, disturbed areas, natural and planted forests, rangelands and grasslands, riparian zones, scrublands and shrublands, urban areas and wetlands. Strawberry guava is a major threat to flora and fauna, especially rare and endemic species, by forming shade-casting thickets with dense mats of surface roots that make it difficult for other species to establish.

Miconia calvescens is a small South American tree initially introduced into Hawaii and Tahiti as an

Morella faya, commonly called the fire tree, is native to the Azores, the Madeira Islands and the Canary

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Rubus ellipticus is a prickly shrub, a wild raspberry, which invades natural forests, agricultural areas, disturbed areas, and rangelands and grasslands. It spreads vegetatively by way of underground shoots and the seeds are dispersed by frugivorous birds and mammals. Hawaii has faced many infestations and this species has displaced the native Hawaiian raspberry (Rubus hawaiiensis).
Native to Argentina, Paraguay and Brazil, Schinus terebinthifolius has invaded Mauritius, Norfolk Island, St. Helena, and the United States (Florida, Hawaii) (Cronk and Fuller, 1995). This tree, commonly called the Brazilian pepper tree, is a successful invader of both disturbed sites and undisturbed natural environments including forests. An aggressive weed, S. terebinthifolius displaces native vegetation and can also cause severe health and respiratory problems for humans (Kendle and Rose, 2001). The African tulip tree, Spathodea campanulata, is an evergreen tree native to West Africa that has been widely planted throughout the tropics and has established in many parts of the Pacific including American Samoa, the Cook Islands, Fiji, Guam, Hawaii and Vanuatu. It invades agricultural areas, forest plantations and natural ecosystems where it smothers trees and crops and eventually displaces them.

Ulex europaeus, or gorse, is a spiny, perennial, evergreen shrub commonly found in disturbed areas, grasslands, shrublands, forest margins and coastal habitats. Once established, it is a very successful and persistent plant that forms dense and impenetrable thickets capable of displacing native plant and tree species. In British Columbia, Canada, for example, gorse represses the regrowth of the commercially important Douglas fir (Pseudotsuga menziesii) (FAO, 2000a).

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Annex 3. International and regional instruments and programmes


Numerous international and regional instruments, binding and nonbinding, have been developed to deal with the problem of alien invasive species. The following have direct or indirect implications for forests and the forest sector.

International
Cartagena Protocol on Biosafety to the Convention on Biological Diversity Montreal, 2000 On 29 January 2000, the Conference of the Parties to the Convention on Biological Diversity adopted a supplementary agreement to the Convention known as the Cartagena Protocol on Biosafety. The Protocol seeks to protect biological diversity from the potential risks posed by living modified organisms resulting from modern biotechnology. It establishes an advance informed agreement (AIA) procedure for ensuring that countries are provided with the information necessary to make informed decisions before agreeing to the import of such organisms into their territory. Links: Cartagena Protocol on Biosafety: www.biodiv.org/biosafety Convention on Biological Diversity (CBD) Nairobi, 1993 One of the key agreements adopted at the 1992 Earth Summit in Rio de Janeiro was the Convention on Biological Diversity which has three main goals: the conservation of biological diversity, the sustainable use of its components, and the fair and equitable sharing of the benefits from the use of genetic resources. Article 8 (h) of the Convention calls on Parties to as far as possible and as appropriate: Prevent the introduction of, control or eradicate those alien species which threaten ecosystems, habitats or species. At the sixth Conference of the Parties (COP-6), Parties adopted guiding principles and a programme of work for the implementation of Article 8 (h) (decision VI/23). Links: Invasive alien species and the CBD: www.biodiv.org/programmes/cross-cutting/alien Article 8 (h): www.biodiv.org/decisions/?dec=VII/13 Guiding principles for the prevention, introduction and mitigation of impacts of invasive alien species: www.biodiv.org/decisions/?dec=VI/23 Convention on the Conservation of Migratory Species of Wild Animals (CMS) Bonn, 1979 The CMS (or Bonn Convention) is an intergovernmental treaty that aims to conserve terrestrial, marine and avian migratory species throughout their range. The Convention states that parties that are Range States of a migratory species listed in Appendix I shall endeavour, that international agreements for the conservation and management of migratory species should aim, to prevent, reduce or control factors that are endangering or are likely to further endanger the species, including strictly controlling the introduction of, or controlling or eliminating, already introduced alien species. Links: CMS: www.cms.int 55

Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES) 1975 CITES is an international agreement that aims to ensure that international trade in specimens of wild animals and plants does not threaten their survival. The Conference of the Parties, at its thirteenth meeting in Bangkok, Thailand in 2004, recommended that Parties: consider the problems of invasive species when developing national legislation and regulations that deal with trade in live animals or plants; consult with the Management Authority of a proposed country of import when considering exports of potentially invasive species, to determine whether there are domestic measures regulating such imports; consider the opportunities for synergy between CITES and the Convention on Biological Diversity, and explore appropriate cooperation and collaboration between the two Conventions on the issue of introductions of alien invasive species. Links: CITES: www.cites.org Trade in alien species - Thirteenth meeting of the Conference of the Parties, Bangkok, Thailand, 2-14 October 2004: www.cites.org/eng/res/13/13-10.shtml Convention on Wetlands of International Importance especially as Waterfowl Habitat (Ramsar Convention) Ramsar, 1971 In Resolution VII.14 on Invasive Species and Wetlands, the seventh Conference of Contracting Parties calls upon parties to address the environmental, economic and social impact of invasive species on wetlands within their jurisdictions and to take account of the methods of control and solutions for combating invasive species. Links: Ramsar Convention: www.ramsar.org Resolution VII.14 COP-7: www.ramsar.org/res/key_res_vii.14e.htm Invasive species and wetlands: www.ramsar.org/cop7/cop7_doc_24_e.htm Food and Agriculture Organization of the United Nations (FAO) Biosecurity is one of FAOs Priority Areas for Inter-disciplinary Action. Biosecurity is a strategic and integrated approach that encompasses the policy and regulatory frameworks that analyse and manage risks in the sectors of food safety, animal life and health, and plant life and health, including associated environmental risk. It covers the introduction of plant pests, animal pests and diseases, and zoonoses, the introduction and release of genetically modified organisms (GMOs) and their products, and the introduction and management of alien invasive species and genotypes. One tool developed by FAO, in association with the organizations responsible for international standard setting in sanitary and phytosanitary (SPS) matters, to assist in providing information to countries to achieve biosecurity is the International Portal on Food Safety, Animal and Plant Health (IPFSAPH). This Web site provides a single access point for the latest versions of international and national standards, regulations and other official materials relating to SPS measures in food and agriculture including fisheries and forestry. Links: FAO Priority Area for Inter-disciplinary Action - Biosecurity for Agriculture and Food Production: www.fao.org/biosecurity FAO Technical Consultation on Biological Risk Management in Food and Agriculture, Bangkok, Thailand, 13-17 January 2003: www.fao.org/es/ESN/food/meetings_biosecurity_en.stm

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Reference document: ftp://ftp.fao.org/es/esn/food/tc_bangkok/tc_brm_03_2en.doc International Portal on Food Safety, Animal and Plant Health (IPFSAPH) - Invasive alien species: www.ipfsaph.org/servlet/CDSServlet?status=ND1jdGh0dHB3d3dmYW9vcmdhb3NpcGZzYXBoaXNzd WVrZXl3b3Jkc2ludmFzaXZlYWxpZW5zcGVjaWVzJjY9ZW4mMzM9KiYzNz1rb3M~ Alien invasive species: impacts on forests and forestry: www.fao.org/forestry/aliens Biosecurity in forestry: www.fao.org/forestry/site/biotec Global Invasive Species Programme (GISP) GISP was established in 1997 to address global threats caused by alien invasive species, and to provide support to the implementation of Article 8(h) of the CBD. In its initial stages, GISP was coordinated by the Scientific Committee on Problems of the Environment (SCOPE) in conjunction with the World Conservation Union (IUCN), CAB International and the United Nations Environment Programme (UNEP). Links: Global Invasive Species Programme: www.gisp.org International Civil Aviation Organization (ICAO) 1998 The ICAO urges its Contracting States to support one another's efforts to reduce the risk of introducing, through civil air transportation, potentially invasive alien species to areas outside their natural range. Links: Resolution A-33-19 - Preventing the introduction of invasive alien species: www.icao.int/icaonet/dcs/9848/9848_en.pdf International Plant Protection Convention (IPPC) The International Plant Protection Convention is an international treaty relating to plant health, to which 132 governments (as of 30 December 2004) currently adhere. The purpose of the IPPC is to secure common and effective action to prevent the spread and introduction of pests of plants and plant products, and to promote appropriate measures for their control. The Convention provides a framework and a forum for international cooperation, harmonization and technical exchange between contracting parties dedicated to these goals. Its implementation involves the collaboration of national plant protection organizations (NPPOs), which are the official services established by governments to discharge the functions specified by the IPPC, and regional plant protection organizations (RPPOs) which function as coordinating bodies on a regional level. The IPPC is governed by the Interim Commission on Phytosanitary Measures (ICPM) which adopts International Standards for Phytosanitary Measures (ISPMs). The ICPM established the IPP as the forum for national reporting and exchange of more general information among the phytosanitary community. The IPPC Secretariat coordinates the activities of the Convention and is hosted by FAO. Links: International Phytosanitary Portal (IPP): www.ippc.int/IPP/En/default.jsp Text of the Convention: www.fao.org/legal/treaties/004t2-e.htm RPPOs: Asia and Pacific Plant Protection Commission (APPPC): www.ippc.int/servlet/CDSServlet?status=ND0xMzQ5NyY2PWVuJjMzPSomMzc9a29z Comunidad Andina (CA): www.ippc.int/servlet/CDSServlet?status=ND0xMzQ5MyY2PWVuJjMzPSomMzc9a29z Comite de Sanidad Vegetal del Cono Sur (COSAVE): www.cosave.org

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Caribbean Plant Protection Commission (CPPC): https://www.ippc.int/servlet/CDSServlet?status=ND0xMzQ3MCY2PWVuJjMzPSomMzc9a29z European and Mediterranean Plant Protection Organization (EPPO): www.eppo.org Inter-African Phytosanitary Council (IAPSC): www.au-appo.org North American Plant Protection Organization (NAPPO): www.nappo.org Organismo Internacional Regional de Sanidad Agropecuaria (OIRSA): ns1.oirsa.org.sv Pacific Plant Protection Organization (PPPO) - Plant Protection Service of the Secretariat for the Pacific Community (SPC): www.spc.int/pps Agreement for the Establishment of the Near East Plant Protection Organization (Rabat, 1993): www.fao.org/legal/treaties/024t-e.htm The agreement for the creation of a 10th RPPO, the Near East Plant Protection Organization (NEPPO), was signed in 1993. Two more countries must deposit their ratification or accession with FAO to bring NEPPO into force. International Union of Forest Research Organizations (IUFRO) Scientific research in IUFRO is spread over a number of divisions which concentrate on particular areas. Division 7 is dedicated to forest health and includes research on all issues relating to the health of forests such as pests, diseases and pollution. Within this division, Unit 7.03.12 exclusively addresses the issue of alien invasive species and international trade Links: IUFRO Unit 7.03.12 - Alien invasive species and international trade: www.iufro.org/science/divisions/division-7 World Conservation Union (IUCN) The Invasive Species Specialist Group (ISSG) is part of the Species Survival Commission (SSC) of IUCN. The ISSG is a global group of 146 scientific and policy experts on invasive species from 41 countries that provides advice on threats from invasives and control or eradication methods to IUCN members, conservation practitioners and policy-makers. IUCN has produced a set of guidelines, the IUCN Guidelines for the Prevention of Biodiversity Loss Caused by Alien Invasive Species, designed to help countries, conservation agencies and concerned individuals to reduce the threats posed by alien invasive species to global biodiversity. Links: IUCN/SSC Invasive Species Specialist Group (ISSG): www.issg.org Global Invasive Species Database: www.issg.org/database IUCN Guidelines for the Prevention of Biodiversity Loss Caused by Alien Invasive Species: www.iucn.org/themes/ssc/pubs/policy/invasivesEng.htm World Health Organization (WHO) Geneva, 1982 - adopted by the 22nd World Health Assembly in 1969, amended by the 26th World Health Assembly in 1973, and the 34th World Health Assembly in 1981 The purpose of the World Health Organizations International Health Regulations is to prevent, protect against, control and provide a public health response to the international spread of disease in ways that are commensurate with and restricted to public health risks, and which avoid unnecessary interference with international traffic and trade. They are intended to strengthen the use of epidemiological principles as applied internationally to: detect, reduce or eliminate the sources from which infection spreads; improve sanitation in and around ports and airports; prevent the vector dissemination; and encourage epidemiological activities on the national level.

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Links: International Health Regulations: www.who.int/csr/ihr/en World Trade Organization (WTO) Article 20 of the General Agreement on Tariffs and Trade (GATT) allows governments to act on trade in order to protect human, animal or plant life or health, provided they do not discriminate or use this as disguised protectionism. In addition, there are two specific WTO agreements dealing with food safety and animal and plant health and safety, and with product standards. The Sanitary and Phytosanitary Measures Agreement (SPS) which allows countries to set their own standards based on science which should be applied only to the extent necessary to protect human, animal or plant life or health. The Technical Barriers to Trade Agreement (TBT) which aims to ensure that regulations, standards, testing and certification procedures do not create unnecessary obstacles. Links: General Agreement on Tariffs and Trade (GATT): www.sice.oas.org/agreemts/wto_e.asp#WTO%20|%20GATT Sanitary and Phytosanitary Measures: www.wto.org/english/tratop_e/sps_e/sps_e.htm

Regional
African Convention on the Conservation of Nature and Natural Resources Algiers, 1968 The contracting States shall undertake to adopt the measures necessary to ensure conservation, utilization and development of soil, water, flora and faunal resources in accordance with scientific principles and with due regard to the best interests of the people. The Convention states that Parties shall take all necessary measures for the protection, conservation, sustainable use and rehabilitation of vegetation cover including, inter alia, taking concrete steps or measures to control fires, forest exploitation, land clearing for cultivation, grazing by domestic and wild animals, and invasive species (Article VIII). Links: Convention text: www.africaunion.org/Official_documents/Treaties_%20Conventions_%20Protocols/nature%20and%20natural%20r ecesource.pdf Agreed Measures for the Conservation of Antarctic Fauna and Flora Brussels, 1964 Participating governments shall prohibit introduction of non-indigenous plants and animals into the Treaty Area except in accordance with a permit. Permits shall be drawn in terms as specific as possible and issued to allow importation only of animals and plants listed in Annex C. When any such animal or plant might cause harmful interference with the natural system if left unsupervised within the Treaty Area, such permits shall require that it be kept under controlled conditions and, after it has served its purpose, it shall be removed from the Treaty Area or destroyed. Governments must undertake to ensure that all reasonable precautions are taken to prevent the accidental introduction of parasites and diseases into the Treaty Area (Article IX (1-4)). Links: Convention text: www.antarctica.ac.uk/About_Antarctica/Treaty/Flora_and_Fauna.html ASEAN Agreement on the Conservation of Nature and Natural Resources

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Kuala Lumpur, 1985 The Agreement states that Parties shall endeavour to regulate and, where necessary, prohibit the introduction of exotic species (Article 3(3c)). Links: Agreement text: www.aseansec.org/1490.htm Convention on the Conservation of European Wildlife and Natural Resources (Bern Convention) Bern, 1979 A legally-binding international instrument in the field of nature conservation, the Bern Convention covers the natural heritage of the European continent and extends to some states of Africa. It aims to conserve wild flora and fauna and their natural habitats and to promote European cooperation in that field. Article 11(2)(b) of the Convention states that Contracting Parties are to strictly control the introduction of non-native species. Links: Convention text: www.coe.int/t/e/Cultural_Cooperation/Environment/Nature_and_biological_diversity/Nature_protection Benelux Convention on Nature Conservation and Landscape Protection Brussels, 1982 The Convention aims at regulating the cooperation between the three Governments of Belgium, Luxembourg and the Netherlands in the field of the conservation, the management and the restoration of nature and landscapes. Parties to the Convention are required to prohibit the introduction of non-native animal species into the wild without authorization from the competent national authority, pre-introduction assessments, and communications between parties about planned introductions (Benelux Council of Ministers Decision 17.10.83). Links: Convention text: sedac.ciesin.org/entri/texts/benelux.landscape.protection.1982.html Convention for the Conservation of the Biodiversity and the Protection of Wilderness Areas in Central America Managua, 1992 This Convention obliges Central American countries to take measures to ensure the conservation and maintenance of biodiversity, and to ensure that its uses are sustainable. Among other provisions, it would obligate countries to identify and protect important areas, both as habitat and for other reasons, and to restore degraded areas. Parties agree that all mechanisms shall be established for the control or eradication of all exotic species which threaten ecosystems, habitats and wild species (Article 24). Links: Convention text: eelink.net/~asilwildlife/ca_bio.html Convention on the Conservation of Nature in the South Pacific Apia, 1976 Parties shall carefully consider the consequences of deliberate introduction into ecosystems of species not previously occurring therein (Article V (4)). Links: Convention text: www.spc.org.nc/coastfish/Asides/conventions/apia.htm North American Free Trade Agreement 60

1982 NAFTA includes a chapter on sanitary and phytosanitary measures which states that parties may adopt, maintain or apply any sanitary or phytosanitary measure necessary for the protection of human, animal, plant life or health in its territory or relating to the introduction, establishment or spread of an animal or plant pest or disease taking into account conditions relating to transportation and handling, between those areas. Links: NAFTA: www.sice.oas.org/trade/nafta.asp Chapter 7 - Section B - Sanitary and Phytosanitary Measures: www.sice.oas.org/trade/nafta/chap-073.asp North American Agreement on Environmental Cooperation (NAAEC) 1993 The Commission for Environmental Cooperation (CEC) is an international organization created by Canada, Mexico and the United States under the North American Agreement on Environmental Cooperation (NAAEC). The CEC was established to address regional environmental concerns, help prevent potential trade and environmental conflicts, and to promote the effective enforcement of environmental law. The Agreement, which complements the environmental provisions of the North American Free Trade Agreement (NAFTA), states that the Council of the CEC may develop recommendations regarding exotic species which may be harmful (Article 10 (2)(h)). Links: Agreement text: www.cec.org/pubs_info_resources/law_treat_agree/naaec Commission for Environmental Cooperation (CEC): www.cec.org Phytosanitary Convention for Africa Kinshasa, 1967 Signed by heads of African States and Governments of the Organization of African Unity (OAU) now the African Union (AU), the Convention calls upon Member States to take such measures of quarantine, certification or inspection, or other necessary measures as may be considered by the Organization of African Unity in respect of any living organisms, plants, plant material, seeds, soil, compost or packing material (including containers) and any other article the importation of which has been considered by the OAU to constitute a threat to agriculture in any part of Africa. Links: Convention text: www.africaunion.org/Official_documents/Treaties_%20Conventions_%20Protocols/Phyto_Sanitary%20Convention _for_Africa.pdf Protocol Concerning Mediterranean Specially Protected Areas Geneva, 1982 The Protocol states that Parties shall take necessary measures regarding the prohibition of the destruction of plant life or animals and of the introduction of exotic species; the regulation of any act likely to harm or disturb the fauna or flora, including the introduction of indigenous zoological or botanical species; and any other measure aimed at safeguarding ecological and biological processes in protected areas (Article 7). Links: Protocol text: sedac.ciesin.org/entri/texts/acrc/mspecp.txt.html Protocol concerning Protected Areas and Wild Fauna and Flora in the Eastern African Region Nairobi, 1985

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The Protocol states that Contracting Parties are to prohibit the intentional or accidental introduction of alien species which may cause significant or harmful changes to the Eastern African region (Article 7) and to regulate any activity likely to harm or disturb the fauna or flora, including the introduction of non-indigenous animal or plant species (Article 10). Links: Convention text: sedac.ciesin.org/entri/texts/EastAfrPro.html Protocol Concerning Specially Protected Areas and Biological Diversity in the Mediterranean Barcelona, 1995 The Protocol calls upon Parties to: regulate the introduction of any species not indigenous to the specially protected area in question or of genetically modified species (Article 6d); regulate the intentional or accidental introduction of non-indigenous or genetically modified species to the wild and prohibit those that may have harmful impacts on the ecosystems, habitats or species in the Protocol area (Article 13(1)); implement all possible measures to eradicate species that have already been introduced when, after scientific assessment, it appears that such species cause or are likely to cause damage to ecosystems, habitats or species in the Protocol area (Article 13(2)). Links: Protocol text: europa.eu.int/eur-lex/pri/en/oj/dat/1999/l_322/l_32219991214en00030017.pdf Protocol for the Implementation of the Alpine Convention in the Field of Nature Protection and Landscape Conservation Chambery, 1994 The Protocol (Article 17) states that Contracting Parties must guarantee that species of wild fauna and flora not native to the region in the recorded past are not introduced. Exceptions are possible when the introduction is needed for specific use and may not have adverse effects for nature and for the landscape. Links: Secretariat of the Alpine Convention: www.convention-alpine.info/alpen.php Protocol on Environmental Protection to the Antarctic Treaty Madrid, 1991 The Protocol (Annex 2, Article 4) states that no species of animal or plant not native to the Antarctic Treaty area shall be introduced onto land or ice shelfs in the Antarctic Treaty area except in accordance with a permit. Permits shall be issued to allow the importation only of the animals and plants listed in Appendix B and shall specify the species, numbers and, if appropriate, age and sex and precautions to be taken to prevent escape or contact with native fauna and flora. Any plant or animal for which a permit has been issued shall, prior to expiration of the permit, be removed from the Antarctic Treaty area or be disposed of by incineration or equally effective means that eliminates risk to native fauna or flora. Any other plant or animal introduced into the Antarctic Treaty area not native to that area, including any progeny, shall be removed or disposed of, by incineration or by equally effective means, so as to be rendered sterile, unless it is determined that they pose no risk to native flora or fauna. The Protocol requires Parties to take precautions to prevent the introduction of micro-organisms (e.g. viruses, bacteria, parasites, yeasts and fungi) not present in the native fauna and flora. Links: Protocol text: www.antarctica.ac.uk/About_Antarctica/Treaty/protocol.html

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