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Mutualisms structure ecosystems and mediate their functioning. They also enhance invasions of many alien species. Invasions disrupt native mutualisms, often leading to population declines, reduced biodiversity, and altered ecosystem functioning. Focusing on three main types of mutualisms (pollination, seed dispersal, and plant-microbial symbioses) and drawing on examples from different ecosystems and from species- and community-level studies, we review the key mechanisms whereby such positive interactions mediate invasions and are in turn influenced by invasions. High interaction generalization is \"the norm\" in most systems, allowing alien species to infiltrate recipient communities. We identify traits that influence invasiveness (e.g., selfing capacity in plants, animal behavioral traits) or invasibility (e.g., partner choice in mycorrhizas/rhizobia) through mutualistic interactions. Mutualistic disruptions due to invasions are pervasive, and subsequent cascading effects are also widespread. Ecological networks provide a useful framework for predicting tipping points for community collapse in response to invasions and other synergistic drivers of global change.

Aim Woody plants were not widely considered to be important invasive alien species until fairly recently. Thousands of species of trees and shrubs have, however, been moved around the world. Many species have spread from planting sites, and some are now among the most widespread and damaging of invasive organisms. This article presents a global list of invasive alien trees and shrubs. It discusses taxonomic biases, geographical patterns, modes of dispersal, reasons for introductions and key issues regarding invasions of non-native woody plants around the world. Location Global. Methods An exhaustive survey was made of regional and national databases and the literature. Correspondence with botanists and ecologists and our own observations in many parts of the world expanded the list. Presence of invasive species was determined for each of 15 broad geographical regions. The main reasons for introduction and dissemination were determined for each species. Results The list comprises 622 species (357 trees, 265 shrubs in 29 plant orders, 78 families, 286 genera). Regions with the largest number of woody invasive alien species are: Australia (183); southern Africa (170); North America (163); Pacific Islands (147); and New Zealand (107). Species introduced for horticulture dominated the list (62% of species: 196 trees and 187 shrubs). The next most important reasons for introduction and dissemination were forestry (13%), food (10%) and agroforestry (7%). Three hundred and twenty-three species (52%) are currently known to be invasive in only one region, and another 126 (20%) occur in only two regions. Only 38 species (6%) are very widespread (invasive in six or more regions). Over 40% of invasive tree species and over 60% of invasive shrub species are bird dispersed. Main conclusions Only between 0.5% and 0.7% of the world's tree and shrub species are currently invasive outside their natural range, but woody plant invasions are rapidly increasing in importance around the world. The objectively compiled list of invasive species presented here provides a snapshot of the current dimensions of the phenomenon and will be useful for screening new introductions for invasive potential.

The literature on biological invasions is biased in favour of invasive species – those that spread and often reach high abundance following introduction by humans. It is, however, also important to understand previous stages in the introduction'naturalization invasion continuum (‘the continuum’), especially the factors that mediate naturalization. The emphasis on invasiveness is partly because most invasions are only recognized once species occupy large adventive ranges or start to spread. Also, many studies lump all alien species, and fail to separate introduced, naturalized and invasive populations and species. These biases impede our ability to elucidate the full suite of drivers of invasion and to predict invasion dynamics, because different factors mediate progression along different sections of the continuum. A better understanding of the determinants of naturalization is important because all naturalized species are potential invaders. Processes leading to naturalization act differently in different regions and global biogeographical patterns of plant invasions result from the interaction of population-biological, macroecological and human-induced factors. We explore what is known about how determinants of naturalization in plants interact at various scales, and how their importance varies along the continuum. Research that is explicitly linked to particular stages of the continuum can generate new information that is appropriate for improving the management of biological invasions if, for example, potentially invasive species are identified before they exert an impact.

Invasion science is a very active subdiscipline of ecology. However, some scientists contend that theoretical integration has been limited and that predictive power remains weak. This paper, focusing on plants, proposes a new multi-pronged research strategy that builds on recent advances in invasion science. More intensive studies on particular model organisms and ecosystems are needed to improve our understanding of the full suite of interacting factors that influence invasions (‘model system research’). At the same time, comparative studies across many study systems are essential for unravelling the context-dependencies of insights that emerge from particular studies (‘multi-site studies’); and quantitative synthesis based on large datasets should be constrained to well-defined theoretical domains (‘focused meta-analysis’). Wealso suggest ways for better integration of information about species biology and ecosystem characteristics (‘invasion syndromes’).Weexpect that a resulting theory of invasions will need to be conceived as a somewhat heterogeneous conglomerate of elements of varying generality and predictive power: laws that apply to well-specified domains, general concepts and theoretical frameworks that can guide thinking in research and management, and in-depth knowledge about the drivers of particular invasions.

The global database of invasive trees and shrubs (Richardson & Rejmánek, 2011; Diversity Distrib. 17, 788-809) has been updated, resulting in a total of 751 species (434 trees and 317 shrubs) from 90 families. Ten originally listed species were deleted (synonyms, inconclusive identification, etc.) and 139 additional invasive species (86 trees and 53 shrubs) are now included in the database. For many species, new records on their adventive distributions are added. The updated database also includes the native ranges for all listed species.

The search for traits associated with plant invasiveness has yielded contradictory results, in part because most previous studies have failed to recognize that different traits are important at different stages along the introduction–naturalization–invasion continuum. Here we show that across six different habitat types in temperate Central Europe, naturalized non-invasive species are functionally similar to native species occurring in the same habitat type, but invasive species are different as they occupy the edge of the plant functional trait space represented in each habitat. This pattern was driven mainly by the greater average height of invasive species. These results suggest that the primary determinant of successful establishment of alien species in resident plant communities is environmental filtering, which is expressed in similar trait distributions. However, to become invasive, established alien species need to be different enough to occupy novel niche space, i.e. the edge of trait space. Plant functional traits may help distinguish introduced species that will become invasive from those that do not. Here, Divíšek et al. show that functional profiles of naturalized plant species are similar to natives, while those of invasive plant species exist at the edge of the functional trait space.

Invasive species cost the global economy billions of dollars each year, but ecologists have struggled to predict the risk of an introduced species naturalizing and invading. Although carefully designed experiments are needed to fully elucidate what makes some species invasive, much can be learned from unintentional experiments involving the introduction of species beyond their native ranges. Here, we assess invasion risk by linking a physiologically based species distribution model with data on the invasive success of 749 Australian acacia and eucalypt tree species that have, over more than a century, been introduced around the world. The model correctly predicts 92% of occurrences observed outside of Australia from an independent dataset. We found that invasiveness is positively associated with the projection of physiological niche volume in geographic space, thereby illustrating that species tolerant of a broader range of environmental conditions are more likely to be invasive. Species achieve this broader tolerance in different ways, meaning that the traits that define invasive success are context-specific. Hence, our study reconciles studies that have failed to identify the traits that define invasive success with the urgent and pragmatic need to predict invasive success.

Aim An increasingly important component of invasive species management involves the formal assessment of risks associated with particular species becoming invasive and causing impact. We evaluated recent developments in risk assessment (RA) for alien species, with special emphasis on species-based pre-border assessments for intentional introductions. Our aim was to identify important advances and key challenges. Location Global. Methods A literature review was done to determine which approaches have been developed and fine-tuned over the last two decades, which of these have worked best and which are most widely used. We identified priorities for improving our ability to assess risks. Results The review is divided into sections on various types and foci of RAs: invasion stage, taxon, ecosystem, assessment method and impact type. RAs for plants are the most advanced, with the Australian Weed Risk Assessment (A-WRA) being the most widely applied and tested protocol. Based on the history of the A-WRA, we highlight advances that have been made in assessing risk of alien species for pre-border control and identify remaining challenges. Main conclusions Currently available RAs have proven to be cost-effective, but there is room for substantial improvement. Further work is needed to separate likelihood and consequence more explicitly, and provide better and more objective means for assessing risks of impact. Types and levels of uncertainty need to be more effectively incorporated. Advanced RA protocols are needed for taxa other than plants and vertebrates. The latest insights from research in invasion ecology need to be incorporated, and advances in other fields must also be taken into account.

Invasive alien species are a major problem for managers of protected areas (PAs) worldwide. Until the 1980s biological invasions were widely considered to be largely confined to anthropogenically disturbed sites and the widespread disruption of ecosystems in PAs by invasive species was not globally perceived as a major threat. A working group of the SCOPE program on biological invasions in the 1980s showed that PAs are not spared from major disruptive effects of invasions. Early research focused on descriptive studies of the extent to which PAs were invaded. More recent research explored drivers of invasion, and in the last decade much work has focused on understanding the impacts of invasions. We review the current understanding of alien plant invasions in PAs, focusing on four themes: (1) the status and macroecological patterns of alien plant invasions; (2) the threats that invasive alien plants (IAPs) pose and the impacts detected to date; (3) the current focus of invasion science in PAs; and (4) research priorities for advancing science-based management and policy. Of a sample of 59 widespread IAP species from a representative sample of 135 PAs globally, trees make up the largest proportion (32%), followed by perennial herbs (17%) and shrubs (15%). About 1857 papers have been published on alien species in PAs; 45% have focused on alien plants. Some textbook examples of impacts by IAPs originate from PAs, illustrating the severe threat to the core function of PAs. Impacts have been quantified at the species and community levels through the displacement and alteration of habitats. In some cases, native species abundance, diversity and estimated species richness have been altered, but reversed following control. At an ecosystem level, invasive plants have radically altered fire regimes in several PAs, in some cases causing regime shifts and transforming woodlands or savannas to grasslands. Invasions have also had a major impact on nutrient cycles. Protected areas are performing an increasingly important part of the global response to stem the rate of environmental change. Despite this, integrated efforts involving science, management and policy that are sufficiently resourced to generate insights on the status and dynamics of IAPs in PAs are insufficient or even lacking. Such efforts are needed to pave the way for monitoring trends, revising legislation and policies, and improving management interventions to reduce the extent and magnitude of impacts of invasive plants in PAs. While policy instruments to support management of non-native species date back to the 1930s, there has been a substantial increase in legislative support and general awareness since the early 2000s. Still, opportunities to improve research for PAs need to be created. Towards this goal, the establishment of a global PA research network could provide a unique vehicle to explore questions across species or functional groups and systems, at a scale currently beyond existing abilities. Developing an integrated global database with standardized, quantitative information could form part of such a networks function.

Species moved by human activities beyond the limits of their native geographic ranges into areas in which they do not naturally occur (termed aliens) can cause a broad range of significant changes to recipient ecosystems; however, their impacts vary greatly across species and the ecosystems into which they are introduced. There is therefore a critical need for a standardised method to evaluate, compare, and eventually predict the magnitudes of these different impacts. Here, we propose a straightforward system for classifying alien species according to the magnitude of their environmental impacts, based on the mechanisms of impact used to code species in the International Union for Conservation of Nature (IUCN) Global Invasive Species Database, which are presented here for the first time. The classification system uses five semi-quantitative scenarios describing impacts under each mechanism to assign species to different levels of impact-ranging from Minimal to Massive-with assignment corresponding to the highest level of deleterious impact associated with any of the mechanisms. The scheme also includes categories for species that are Not Evaluated, have No Alien Population, or are Data Deficient, and a method for assigning uncertainty to all the classifications. We show how this classification system is applicable at different levels of ecological complexity and different spatial and temporal scales, and embraces existing impact metrics. In fact, the scheme is analogous to the already widely adopted and accepted Red List approach to categorising extinction risk, and so could conceivably be readily integrated with existing practices and policies in many regions.