Scientists' warning on invasive alien species

Scientists’warning on invasive alien species Petr Pyšek 1,2,3 *, Philip E. Hulme 4 , Dan Simberloff 5 , Sven Bacher 6 , Tim M. Blackburn 7,8,3 , James T. Carlton 9 , Wayne Dawson 10 , Franz Essl 11,3 , Llewellyn C. Foxcroft 3,12 , Piero Genovesi 13,3 , Jonathan M. Jeschke 14,15,16 , Ingolf Kühn 17,18,19 , Andrew M. Liebhold 20,21 , Nicholas E. Mandrak 22 , Laura A. Meyerson 23 , Aníbal Pauchard 24,25 , Jan Pergl 1 , Helen E. Roy 26 , Hanno Seebens 27 , Mark van Kleunen 28,29 , Montserrat Vilà 30,31 , Michael J. Wingfield 32 and David M. Richardson 3 1 Czech Academy of Sciences, Institute of Botany, Department of Invasion Ecology, Pru˚honice, CZ-252 43, Czech Republic 2 Department of Ecology, Faculty of Science, Charles University, Vinicˇná 7, Prague, CZ-128 44, Czech Republic 3 Centre for Invasion Biology, Department of Botany & Zoology, Stellenbosch University, Matieland, 7602, South Africa 4 Bio-Protection Research Centre, Lincoln University, Canterbury, New Zealand 5 Department of Ecology and Evolutionary Biology, University of Tennessee, Knoxville, TN, U.S.A. 6 Department of Biology, University of Fribourg, Fribourg, Switzerland 7 Centre for Biodiversity and Environment Research, Department of Genetics, Evolution, and Environment, University College London, London, WC1E 6BT, U.K. 8 Institute of Zoology, Zoological Society of London, Regent’s Park, London, NW1 4RY, U.K. 9 Maritime Studies Program, Williams College –Mystic Seaport, 75 Greenmanville, Mystic, CT, 06355, U.S.A. 10 Department of Biosciences, Durham University, South Road, Durham, DH1 3LE, U.K. 11 Division of Conservation Biology, Vegetation and Landscape Ecology, Department of Botany and Biodiversity Research, University of Vienna, Vienna, Austria 12 Conservation Services, South African National Parks, Private Bag X402, Skukuza, 1350, South Africa 13 ISPRA, Institute for Environmental Protection and Research and Chair IUCN SSC Invasive Species Specialist Group, Rome, Italy 14 Leibniz-Institute of Freshwater Ecology and Inland Fisheries (IGB), Müggelseedamm 310, Berlin, 12587, Germany 15 Institute of Biology, Freie Universität Berlin, Königin-Luise-Str. 1-3, Berlin, 14195, Germany 16 Berlin-Brandenburg Institute of Advanced Biodiversity Research (BBIB), Königin-Luise-Str. 2-4, Berlin, 14195, Germany 17 Department Community Ecology, Helmholtz Centre for Environmental Research –UFZ, Theodor-Lieser-Str. 4, Halle, 06120, Germany 18 Geobotany & Botanical Garden, Martin Luther University Halle-Wittenberg, Am Kirchtor 1, Halle, 06108, Germany 19 German Centre for Integrative Biodiversity Research (iDiv) Halle-Jena-Leipzig, Deutscher Platz 5e, Leipzig, 04103, Germany 20 US Forest Service Northern Research Station, 180 Canfield St., Morgantown, West Virginia, U.S.A. 21 Faculty of Forestry and Wood Sciences, Czech University of Life Sciences Prague, Prague, CZ-165 00, Czech Republic 22 Department of Biological Sciences, University of Toronto, 1265 Military Trail, Toronto, Ontario, M1C 1A4, Canada 23 Department of Natural Resources Science, The University of Rhode Island, Kingston, Rhode Island, 02881, U.S.A. 24 Facultad de Ciencias Forestales, Universidad de Concepción, Concepción, Chile 25 Institute of Ecology and Biodiversity, Santiago, Chile 26 U.K. Centre for Ecology & Hydrology, Wallingford, OX10 8BB, U.K. 27 Senckenberg Biodiversity and Climate Research Centre (SBiK-F), Senckenberganlage 25, Frankfurt am Main, 60325, Germany 28 Ecology, Department of Biology, University of Konstanz, Universitätsstrasse 10, Constance, 78457, Germany 29 Zhejiang Provincial Key Laboratory of Plant Evolutionary Ecology and Conservation, Taizhou University, Taizhou, 318000, China 30 Estación Biológica de Doñana (EBD-CSIC), Avd. Américo Vespucio 26, Isla de la Cartuja, Sevilla, 41092, Spain 31 Department of Plant Biology and Ecology, University of Sevilla, Sevilla, Spain 32 Forestry and Agricultural Biotechnology Institute (FABI), University of Pretoria, Pretoria, South Africa *Address for correspondence (Tel: +420 271015266; E-mail: [email protected]). Biological Reviews 95 (2020) 1511–1534 © 2020 The Authors. Biological Reviews published by John Wiley & Sons Ltd on behalf of Cambridge Philosophical Society. This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited. Biol. Rev. (2020), 95, pp. 1511–1534. 1511 doi: 10.1111/brv.12627 ABSTRACT Biological invasions are a global consequence of an increasingly connected world and the rise in human population size. The numbers of invasive alien species –the subset of alien species that spread widely in areas where they are not native, affecting the environment or human livelihoods –are increasing. Synergies with other global changes are exacerbating current invasions and facilitating new ones, thereby escalating the extent and impacts of invaders. Invasions have complex and often immense long-term direct and indirect impacts. In many cases, such impacts become apparent or problematic only when invaders are well established and have large ranges. Invasive alien species break down biogeographic realms, affect native species richness and abundance, increase the risk of native species extinction, affect the genetic composition of native populations, change native animal behaviour, alter phylogenetic diversity across communities, and modify trophic networks. Many invasive alien species also change ecosystem functioning and the delivery of ecosystem services by altering nutrient and contaminant cycling, hydrology, habitat structure, and disturbance regimes. These biodiversity and ecosystem impacts are accelerating and will increase further in the future. Scientific evidence has identified policy strategies to reduce future invasions, but these strategies are often insufficiently implemented. For some nations, notably Australia and New Zealand, biosecurity has become a national priority. There have been long-term successes, such as eradication of rats and cats on increasingly large islands and biological control of weeds across continental areas. However, in many countries, invasions receive little attention. Improved international cooperation is crucial to reduce the impacts of invasive alien species on biodiversity, ecosystem services, and human livelihoods. Countries can strengthen their biosecurity regulations to implement and enforce more effective management strategies that should also address other global changes that interact with invasions. Key words: biological invasions, biosecurity, global change, environmental impacts, invasion dynamics, invasion hotspots, naturalization, policy, protected areas, socioeconomic impacts CONTENTS I. Introduction ......................................................................1513 (1) Relevance to Scientists’warning initiative .............................................1513 (2) What is an invasive alien species? ...................................................1513 (3) Aims and scope of the paper .......................................................1513 II. Where do we stand? The state of biological invasions .......................................1513 (1) Global extent of invasions .........................................................1513 (2) Introduction pathways ............................................................1515 (3) Driving factors ..................................................................1516 (4) Dynamics of invasions ............................................................1517 (5) Invasions in protected areas ........................................................1517 III. Why should we care? The impacts of biological invasions ....................................1518 (1) Environmental impacts ...........................................................1518 (2) Impacts on human well-being and livelihoods ..........................................1520 IV. What tools do we have? Instruments, regulations and management ............................1520 (1) International agreements, legislation, and voluntary self-regulation .........................1520 (2) National biosecurity programs ......................................................1523 (3) Technological advances in management: from classical control to gene editing ................1523 (4) Surveillance and monitoring: the key role of citizen science ...............................1524 V. Invasions in the future: what’s next? ....................................................1524 VI. Research priorities ..................................................................1525 (1) Invasions require both local- and global-scale solutions ...................................1525 (2) Management interventions need to be objectively prioritized ..............................1526 (3) Protected areas need special attention ................................................1526 (4) More effective protocols are needed for engaging with the public and societal actors ............1526 (5) Forecasting and scenario development must give more attention to synergies of invasions with climate change and other environmental changes ................................................1526 VII. Conclusions .......................................................................1526 VIII. Acknowledgements .................................................................1527 IX. References ........................................................................1527 Biological Reviews 95 (2020) 1511–1534 © 2020 The Authors. Biological Reviews published by John Wiley & Sons Ltd on behalf of Cambridge Philosophical Society. 1512 Petr Pyšek et al. I. INTRODUCTION (1) Relevance to Scientists’warning initiative Nearly three decades ago, a community of eminent scientists warned that humans were on a collision course with the natural world. They cited concerns regarding ozone depletion, freshwater availability, marine life depletion, ocean dead zones, forest loss, biodiversity destruction, climate change, and continued human population growth (Union of Concerned Scientists, 1992). Twenty-five years later, Ripple et al. (2017) evaluated the human response based on their analysis of time-series data and concluded that humanity had failed to make sufficient progress over that period in dealing with the environmental challenges. Indeed, they concluded that most of these problems had worsened. The original 1992 call was supported by more than 1,700 scientists, while 25 years later over 15,000 scientists added their signatures to the recent declaration (Ripple et al., 2017). Comparing the two documents reveals an important difference in focus as regards biodiversity loss and species extinctions. With respect to biodiversity, the 1992 warning explicitly highlighted deforestation, species loss, and climate change but did not mention invasive alien species (IAS). However, in the second call, besides stressing the need to respond to indirect drivers of biodiversity loss (e.g. to limit population growth, reassess the role of an economy rooted in growth, or reduce greenhouse gas emissions), Ripple et al. (2017) addressed options available to alter biodiversity decline, such as protecting and restoring ecosystems, halting defaunation, and constraining the spread of IAS. Since 1992, the importance of taking action against IAS globally has been widely recognized (Millennium Ecosystem Assessment, 2005). IAS are, for example, listed among the major indicators of global biodiversity decline (Butchart et al., 2010). The recent global assessment report on biodiversity and ecosystem services by the Intergovernmental Science-Policy Platform on Biodiversity and Ecosystem Services (IPBES) ranked IAS fifth among direct drivers of change in nature with the largest relative global impacts, after changes in land and sea use, direct exploitation of organisms, climate change, and pollution (Brondizio et al., 2019). As the next step, the IPBES has initiated a global assessment on IAS that will also address management and policy needs and is expected to deliver results by 2023. (2) What is an invasive alien species? Alien species (as opposed to native species) are those whose presence in a region is attributable to human actions, deliberate or inadvertent, that enabled them to overcome biogeographical barriers (Richardson et al., 2000; Pyšek et al., 2004; Richardson, Pyšek, & Carlton, 2011; Essl et al., 2018). Some alien species become established (i.e. they reproduce regularly to form self-replacing populations); a subset of these spread rapidly over substantial distances from introduction sites, a process that forms the basis for the definition of invasive species (Richardson et al., 2000; Occhipinti-Ambrogi & Galil, 2004; Pyšek et al., 2004; Blackburn et al., 2011). Another definition, supported by the International Union for Conservation of Nature (IUCN), the Convention on Biological Diversity, and the World Trade Organization, classifies as ‘invasive’only those alien species that have a harmful effect on the economy, environment, or health (IUCN, 2000). (3) Aims and scope of the paper To support global initiatives addressing the loss of biodiversity and ecosystem services, this paper presents a comprehensive global overview of a major environmental change – invasion by alien species. We (i) appraise the current state of biological invasions in marine, freshwater, and terrestrial ecosystems; (ii) show that current societal responses are insufficient to address impacts of IAS on ecosystems, biodiversity, and human well-being and to mitigate future risks; and (iii) argue that a warning to humanity regarding the threats posed by IAS is both timely and relevant to complement other focused papers pointing to current threats to nature and the importance of nature for humans (e.g. Finlayson et al., 2019; Mammola et al., 2019; Cardoso et al., 2020). Given the recent rise and intensity of research on IAS at a global scale (e.g. Early et al., 2016; Paini et al., 2016; Dawson et al., 2017; Seebens et al., 2017), we provide a detailed update on the extent of invasions and their impacts worldwide, identify drivers that promote invasions, and explore how invasions interact with other biodiversity stressors and global changes. We emphasize that as our knowledge increases the problems associated with invasions are becoming clearer and require urgent increased attention, and that policy makers and the public should prioritize actions to stem invasions and their impacts. We address four questions that must be answered as the basis for mitigating problems associated with biological invasions. (i) Where do we stand? (ii) Why should we care? (iii) What tools are available to deal with these problems? (iv) What comes next? We treat these questions separately and then provide recommendations for policy, management, and research. II. WHERE DO WE STAND? THE STATE OF BIOLOGICAL INVASIONS (1) Global extent of invasions The availability and accessibility of global data on alien organisms and their distribution have improved greatly over the last few decades. Comprehensive accounts are now available on established and/or invasive alien species of vascular plants (van Kleunen et al., 2015, 2019; Pyšek et al., 2017b), bryophytes (Essl et al., 2015), terrestrial snails (Capinha et al., 2015), ants, spiders [see Dawson et al., 2017 and references therein], fishes (Tedesco et al., 2017), amphibians (Capinha et al., 2017), reptiles (Kraus, 2009, 2015; Capinha Biological Reviews 95 (2020) 1511–1534 © 2020 The Authors. Biological Reviews published by John Wiley & Sons Ltd on behalf of Cambridge Philosophical Society. Global overview of biological invasions 1513 et al., 2017), birds (Blackburn, Cassey, & Lockwood, 2008; Blackburn, Lockwood, & Cassey, 2009; Dyer et al., 2017; Dyer, Redding, & Blackburn, 2017), and mammals (Long, 2003; Dawson et al., 2017), and many of the global hotspots of established alien species across taxa have been identified (Fig. 1). Additional comprehensive accounts are available for many alien taxa at continental, regional, or national scales. These databases and analyses of ecological patterns and impacts associated with alien species have resulted from large international collaborations and rapid technological developments including data-sharing and analysis tools. The IUCN SSC Invasive Species Specialist Group maintains two global databases: the Global Invasive Species Database (www.iucngisd.org), which contains profiles of key IAS, and the Global Register of Introduced and Invasive Alien Species (www.griis.org; Pagad et al., 2018), which was developed with a mandate of the Convention on Biological Diversity (CBD) and collates data on alien species in all taxonomic groups for all nations (Pagad et al., 2015, 2018). Projects such as DAISIE [Delivering Alien Species Inventories for Europe (DAISIE, 2009; Hulme et al., 2009)], USGS reports on alien species (Fuller & Neilson, 2015; Simpson & Eyler, 2018), NOBANIS (North European and Baltic Network on Invasive Alien Species; www.nobanis.org), and NEMESIS (https://invasions.si.edu/nemesis) exemplify cross-taxonomic initiatives that have produced such databases for particular regions (Hulme & Weser, 2011). The CABI Invasive Species Compendium is another detailed source of information. The World Register of Introduced Marine Species (WRiMS) records many of the marine species included in the World Register of Marine species (WRoMS) that have been introduced deliberately or accidentally (Pagad et al., 2018; Ahyong et al., 2019). The FAO Database on Introductions of Aquatic Species (DIAS) contains mostly data on the distribution of alien freshwater taxa, particularly fishes, molluscs, and crustaceans (FAO, 2019). Detailed critical assessments also exist of the introduction status of several groups, for example trees and shrubs (Rejmánek & Richardson, 2013). However, for some important groups of organisms, particularly many invertebrates and microorganisms, data on alien species distributions are still very limited (Fisher et al., 2012; Thakur et al., 2019), and regional biases also exist in terms of data availability associated with socioeconomic status and development (Pyšek et al., 2008; Nuñez & Pauchard, 2010). Through these efforts, we now have a good knowledge of the numbers of established and –to a lesser extent –invasive alien species for many taxonomic groups in various regions and estimates of their total numbers globally. For vascular plants, the most recent figures report 14,000 species with established alien populations in at least one region, constituting 4% of the world flora. North America and Europe have accumulated the largest numbers of established alien species. Fig 1. Hotspots and coldspots of cross-taxon established alien species richness across eight taxonomic groups: vascular plants, ants, spiders, freshwater fishes, amphibians, reptiles, birds, and mammals, calculated as in Dawson et al. (2017). Cross-taxon values were calculated as averages of established alien species richness per taxonomic group (scaled according to the maximum value) in a region with data available. Only TDWG level-4 regions (countries, federal states and islands/archipelagos) that were modelled by Dawson et al. (2017) were included (N=439). Cross-taxon established alien species richness of grey-bordered regions was calculated from three or fewer taxonomic groups, and of black-bordered regions from four or more taxonomic groups. Crosstaxon established alien species richness is displayed in percentile categories; upper and lower 2.5% and 10% regions are indicated separately from the remaining upper and lower 50% regions. Regions filled in grey lacked information on established alien species (Antarctica was excluded from the analysis). Biological Reviews 95 (2020) 1511–1534 © 2020 The Authors. Biological Reviews published by John Wiley & Sons Ltd on behalf of Cambridge Philosophical Society. 1514 Petr Pyšek et al. Continents in the Northern Hemisphere have been the major donors of established alien plant species to other continents, and biomes in the New World and in temperate and mediterranean-type climates are generally more invaded than those in arid and warm climates (van Kleunen et al., 2015, 2019; Pyšek et al., 2017b). Estimates of the total number of invasive alien plant species suggest that 2500 species have achieved this status (Pagad et al., 2015); South Africa, India, California, Cuba, Florida, Queensland, and Japan are regions with the highest numbers of reported invasive plant species (Pyšek et al., 2017b). For invertebrates, a global database of alien species richness of terrestrial gastropods documents that at least 175 species have become established across 56 countries. These data show that human-mediated dispersal has broken down biogeographic barriers defined by native species distributions (Capinha et al., 2015). Most crayfish species in Europe today are alien (10 established alien species versus five natives), and the aliens also reach much higher abundances (Kouba, Petrusek, & Kozák, 2014). Alien insect species outnumber invasions of all other animal taxa, with North America having the greatest number of non-native insects (3200 species; Liebhold et al., 2018). Microbes including animal and plant pathogens are arguably the most poorly documented of all IAS. This is largely due to their small size, taxonomic challenges, and difficulties in determining whether taxa are native or alien in particular environments (Fisher et al., 2012; Cowan et al., 2013; Crous, Hawksworth, & Wingfield, 2015; Thakur et al., 2019). Similarly, alien fungi are insufficiently studied (Roy et al., 2017). Invasions by vertebrates are relatively well documented. A global database of freshwater fish distributions in 3,119 drainage basins shows that 8,128 inter-basin introductions of 745 alien species (of 15,000 freshwater fish species globally) have led to established alien populations (Tedesco et al., 2017). The Colorado (100 species) and Mississippi (73 species) basins have the most established alien fish species. The greatest proportions of established alien fish species are found in temperate regions of Europe, North America, and South America (FAO, 2019). For birds, currently the beststudied group of vertebrate invaders, Dyer et al. (2017) collated data on more than 3,660 dated introductions, involving 971 species, from 1500 to 2000; 37% of these species have become established. Notable hotspots of alien bird species richness are the United States (including the Hawaiian Islands), the Caribbean, UK, Japan, Taiwan, Hong Kong, New Zealand, Australia, Persian Gulf States, and the Mascarene Islands, largely driven by spatial patterns of deliberate population introduction (Dyer et al., 2017). A global study of other vertebrate groups documented 78 and 198 alien amphibian and reptile species, respectively, established in at least one of the 359 world regions considered; invasions of herptiles are accelerating rapidly, particularly on islands (Capinha et al., 2017). Globally, a recent analysis revealed that islands and coastal mainland regions are hotspots of established alien species richness across multiple taxonomic groups (Dawson et al., 2017; Fig. 1). Regions with high per capita gross domestic product, high human population densities, and large surface areas support the most established alien species, probably because these characteristics are positively related to the numbers of alien species introduced. The three regions with the most established alien species, after accounting for area, are the Hawaiian Islands, New Zeal- and’s North Island, and the Lesser Sunda Islands of Indonesia. The Hawaiian Islands, long known for devastating environmental impacts of biological invasions (Vitousek, Loope, & Stone, 1987), harbour many alien species in all taxonomic groups considered, including in the marine environment (Carlton & Eldredge, 2015). Almost half of New Zealand’sflora consists of alien plants (Essl et al., 2019a; Hulme, 2020), and predatory mammals have been a major problem for naïve native bird species that evolved without native mammal predators. Florida is the top hotspot for alien species among continental regions, with the Burmese python (Python bivittatus) as a well-known reptile example (Dorcas et al., 2012). These global patterns can tell us which regions have the greatest numbers of naturalized and invasive alien species, but they do not tell us how the management burden they impose can be reduced. This requires us to consider, amongst other things, how species have been introduced. (2) Introduction pathways The Strategic Plan for Biodiversity (2011–2020) of the CBD calls for urgent action by the Parties (i.e. signatory States) to identify and prioritize alien species pathways and to implement measures to manage pathways to prevent alien species introduction and establishment (CBD, 2014). Six broad mechanisms by which alien species might be introduced to a region have been described (Hulme et al., 2008): deliberate release (e.g. game animals, sport fishes, pets); escape from captivity (e.g. ornamental garden plants, pets); contaminants of commodities (e.g. weed seeds, pest insects, microbial pathogens); stowaways on transport vectors (e.g. marine organisms fouling ship hulls or in ballast water, latent endophytic pathogens in plants); via anthropogenic corridors (such as through the Suez and Panama Canals); or unaided spread from other invaded regions. The intentional pathways ‘escape’and ‘release’are most important for plants and vertebrates, whereas for invertebrates, algae, fungi, and microorganisms, unintentional ‘contaminant’and ‘stowaway’ transport pathways prevail; representation of these pathways differs only slightly among marine, freshwater, and terrestrial environments (Saul et al., 2017). On average, IAS with the highest impacts are associated with multiple pathways (Pergl et al., 2017). Historically, many species were deliberately released for economic, recreational, or aesthetic benefits (Lever, 1992). Although authorities are much more cautious today about such releases and generally impose stricter controls on introductions than previously, there are new challenges. For example, some conservationists advocate translocating Biological Reviews 95 (2020) 1511–1534 © 2020 The Authors. Biological Reviews published by John Wiley & Sons Ltd on behalf of Cambridge Philosophical Society. Global overview of biological invasions 1515 individuals of certain species threatened by climate change to new regions predicted to favour population persistence. This strategy, termed managed relocation or assisted migration/ colonization, often involves moving species to sites where they are not currently found and may never have been native (Loss, Terwilliger, & Peterson, 2011). This approach can potentially launch invasions (Ricciardi & Simberloff, 2009), and plans to undertake such movements must be carefully assessed (Richardson et al., 2009), because such relocations are in their infancy and lack evidence that they will achieve their goals. However, from historic intentional introductions of biocontrol agents, several examples of unexpected nontarget effects are famous (e.g. cane toad Bufo marinus; Shanmuganathan et al., 2010). Ornamental plants have escaped from gardens for centuries (Hanspach et al., 2008), and ornamental horticulture continues to be a major driver of alien plant invasions (van Kleunen et al., 2018), even in protected areas (Foxcroft, Richardson, & Wilson, 2008). The dramatic recent growth in trade of unusual pets is another growing threat (Lockwood et al., 2019). Europe alone contains an estimated 54 million individual ornamental birds, 28 million small mammals, 14 million aquaria fishes, and nine million reptiles owned as pets; many of these species can establish outside of captivity, especially under future climate scenarios (Hulme, 2015). These pets might also be important vectors of animal and human diseases, particularly those pets sourced from the wild (Day, 2011). Despite such threats, movement of endo- and ectoparasitic contaminants remains largely unregulated (Hulme, 2014b) and becomes even more difficult to manage effectively in an era of a rapidly growing ‘bioweb’of online commerce of living species (Carlton, 2011). Global shipping expanded enormously after World War II and is projected to increase rapidly in the coming decades (Sardain, Sardain, & Leung, 2019). Consequently, many thousands of species may be transported around the world as stowaways in ballast water (Carlton & Geller, 1993) and as contaminants of transported goods to regions that are becoming increasingly susceptible to new invasions owing to climate warming. Marine invasions are also being exacerbated by the dramatic increase in use of non-biodegradable plastics since the second half of the 20th century, depositing billions of tons of plastics globally at the land-sea interface. A new mechanism for ocean rafting is created when these plastics are swept into the ocean by tsunamis or by the increasing (owing to climate change; Peduzzi et al., 2012) number and size of cyclonic storms (hurricanes, monsoons, typhoons). Whereas biodegradable trees, root masses, seeds, and other ephemeral materials such as pumice facilitated natural dispersal of species across oceans for millions of years, plastics create rafts that can last for decades, permitting more species to be transported as passengers far longer and further (Carlton et al., 2017). Canals have been instrumental in linking once-separated biogeographical regions and facilitating the spread of IAS, with many such corridors expanded to permit larger vessels (Hulme, 2015) and new ones proposed for construction (e.g. the Nicaragua Canal; Huete-Perez, Meyer, & Avarez, 2015). A worrying new global corridor has emerged since the end of the 20th century –the permanent opening of the Arctic Ocean is increasingly permitting the flow of species (presumably both marine and terrestrial) between the Atlantic and Pacific Oceans. As with ocean plastic rafting, we have yet to fully grasp the short- and long-term consequences of the disappearance of the ice-bound Arctic, which has long formed an impassable barrier between oceans and continents (Ricciardi et al., 2017). The dissolution of this northern ice cap now opens a huge corridor, not only for species moving north and for species moving between the Atlantic and Pacific Oceans by ocean currents, but for new fleets of exploratory, cargo, fishing, and tourist vessels, which will inadvertently transport marine and terrestrial species (Ricciardi et al., 2017; Chan et al., 2018). By contrast, the Antarctic is land surrounded by ocean, with a circumpolar current that, while long isolating the continent, may now be bridged by increasing climate-induced storm-driven dispersal (Fraser et al., 2018; Avila et al., 2020). Antarctica has been described as the “final frontier for marine biological invasions”(McCarthy et al., 2019, p. 2221), with formerly ice-bound shores now available for colonization by poleward-moving species. Invasions in Antarctica are being accelerated by new facilities and new forms of tourism accompanied by increased ship traffic, such that more alien species have already been observed (Huiskes et al., 2014; McGeoch et al., 2015; Cardenas et al., 2020) and even more are predicted (Duffy et al., 2017; Hughes et al., 2020). Many terrestrial species are also accidentally transported in trade. For example, wood packaging material often harbours bark and wood-boring insects and microbes; recent increases in trade have produced an explosion of tree-killing insects and pathogens introduced to new regions (Aukema et al., 2010; Paap et al., 2018), with large impacts on forests (Seidl et al., 2018; Fei et al., 2019). Pathogens have been moved with apparently healthy plant germplasm as part of the natural endophytic microbiome, only to emerge as aggressive plant pathogens in new environments. This pathway of introduction has only recently been recognized through the emergence of metagenomic technologies and is particularly relevant for microbial invasives (Marsberg et al., 2017). Invasive alien species are also increasingly spreading without direct human assistance from one region where they have been introduced to other regions. Examples include the ruddy duck (Oxyura jamaicensis)migrating from the UK to Spain, or the currant-lettuce aphid (Nasonovia ribisnigri), dispersing on wind currents from New Zealand to Tasmania and subsequently throughout Australia. This unaided pathway poses major challenges for international regulation as well as biosecurity measures within individual countries (Hulme, 2015). (3) Driving factors The number, rate, and magnitude of biological invasions are shaped by both direct and indirect drivers. Direct drivers of Biological Reviews 95 (2020) 1511–1534 © 2020 The Authors. Biological Reviews published by John Wiley & Sons Ltd on behalf of Cambridge Philosophical Society. 1516 Petr Pyšek et al. invasion can be both natural and anthropogenic and directly affect species physiology, behaviour, and/or demography. Among the best-studied direct drivers are climate change (Walther et al., 2009), land-use change providing new habitats (Chytrý et al., 2008, 2009), pollution (Crooks, Chang, & Ruiz, 2011), and the facilitative effect of other alien species through a process termed invasional meltdown (Simberloff & Von Holle, 1999; Braga et al., 2018; Redding et al., 2019). Indirect drivers mostly operate diffusely by altering and influencing direct drivers, as well as other indirect drivers. They do not impact alien species directly but instead do so by affecting the level, direction, or rate of direct drivers. Global indirect drivers include economic, demographic, governance, technological, and cultural processes. For example, a well-known correlate of alien species richness is economic activity, frequently measured by gross domestic product (Hulme, 2009; Pyšek et al., 2010). Economic activity acts directly by increasing probabilities of species introductions (Hanspach et al., 2008; Maurel et al., 2016; Dyer et al., 2017) or indirectly through other variables, such as the movement of particular commodities, eutrophication, or the intensity of anthropogenic disturbance (Pyšek et al., 2010). By contrast, issues such as governance, culture, or the role of institutions as indirect drivers of biological invasions have been understudied. This may be an important oversight that substantially impedes our understanding of alien species introductions. Invasive alien species often drive change, but they can also be passengers of other human-caused alterations, such as habitat degradation or climate change, that promote colonization and invasion (Didham et al., 2005; MacDougall & Turkington, 2005). Urban habitats are well documented as hotspots of alien plant species establishment and spread because of high colonization and propagule pressures resulting from trade, traffic, horticulture, and frequent and intense disturbances (Hulme, 2009; Kühn, Wolf, & Schneider, 2017). Although studies on the role of human-induced disturbance in animal invasions are less conclusive (Nordheimer & Jeschke, 2018), such research provides a strong signal that biological invasions should be considered together with other global changes. (4) Dynamics of invasions The acceleration of alien species introductions and invasions has been highlighted for several regions (Hulme et al., 2009). The recent IPBES biodiversity and ecosystem services global assessment estimates that numbers of IAS per country have risen by about 70% since 1970 across the 21 countries with detailed records (Brondizio et al., 2019). The most robust analysis of long-term temporal trends in biological invasions is based on more than 45,000 first records of over 16,000 alien species that became established following introduction (Seebens et al., 2017). These data facilitated an analysis of the accumulation of alien species over long periods, showing that for all groups of organisms on all continents, the numbers of alien species have increased continuously over the last 200 years (Fig. 2). Indeed, for most taxonomic groups, rates of first recorded introductions are higher now than at any other time, no signs of a slow-down are evident, and many new invasions will be discovered in the near future given the typical time lags between introductions, establishment, and spread (Crooks, 2005; Jeschke & Strayer, 2005). Because 37% of all recorded alien species have become established recently, between 1970 and 2014 (Seebens et al., 2017), we can expect many more cases of establishment in the future if new arrivals are left unchecked. The growing number of alien species introductions and their subsequent establishment highlights the urgent need for more effective measures for prevention, early detection, and control of IAS (Seebens et al., 2017). Even after many centuries of invasions, the rate of emergence of new alien species is still high: as many as a quarter of first records for the period 2000–2005 were of species not previously recorded as alien species anywhere in the world. These emerging alien species have no invasion history; their potential spread and impacts will therefore be difficult to predict (Seebens et al., 2018). For this reason, and because the extent and magnitude of other global-change factors are changing rapidly, predicting future invasions and their impacts based on the dynamics of historical invasions is likely to lead to a substantial underestimate. (5) Invasions in protected areas Protected areas cover 13% of the terrestrial world (Jenkins & Joppa, 2009), with freshwater often included among the figure for terrestrial areas (Juffe-Bignoli et al., 2016), and 7.7% of the ocean (www.protectedplanet.net); they are a key component of the societal response to environmental degradation (Gaston et al., 2008; Conroy et al., 2011). However, protected areas remain vulnerable to invasions: they suffer from impacts at the species and community levels, through the alteration of habitats, regime shifts, and through diverse undesired effects on native species abundance, diversity, and richness (Foxcroft et al., 2013; Hulme et al., 2014). Few protected areas are completely free of alien plants (Foxcroft et al., 2017), and alien plants can invade natural areas that have not experienced obvious anthropogenic disturbances, such as the Gros Morne National Park in boreal Canada (Rose & Hermanutz, 2004). In 2007, a Global Invasive Species Program (GISP) report identified 487 protected areas globally where invasive alien plants threaten biodiversity (De Poorter, 2007). At the continental scale, the problem is also accelerating. For example, as early as the 1980s, alien plants and animals were perceived as threatening natural resources in 300 areas managed by the USA National Park Service (Houston & Schreiner, 1995). Invasive plants are almost universally regarded as a major threat by managers of protected areas (Goodman, 2003; Randall, 2011; Pyšek et al., 2013). Even in high-elevation protected areas, in isolated mountain landscapes, invasive alien species have become a problem (Alexander et al., 2016). These trends have sometimes been reversed following the implementation of control efforts (Simberloff etal., 2011,2018; Biological Reviews 95 (2020) 1511–1534 © 2020 The Authors. Biological Reviews published by John Wiley & Sons Ltd on behalf of Cambridge Philosophical Society. Global overview of biological invasions 1517 Shackleton et al., 2020), and several studies have shown that protected area boundaries provide some resistance to colonization by alien plants (Lonsdale, 1999; Pyšek, Jarošík, & Kucˇera, 2003; Foxcroft et al., 2011). However, as human populations adjacent to many protected areas are growing rapidly (Wittemyer et al., 2008), colonization and propagule pressures will increase. More research is needed to strengthen management actions (Foxcroft et al., 2017), because, for example, few assessments or studies on the impacts of invasions in protected areas provide management recommendations (Genovesi & Monaco, 2013; Hulme et al., 2014). For plants, a global assessment showed that 37% of 282 quantitative studies on impacts of IAS in the peer-reviewed literature originated from research in protected areas. However, geographical biases are evident –much more research has been conducted in the Americas and on Pacific Islands than in Africa, Asia, and Europe (Hulme et al., 2014). A fundamental problem is that current approaches for estimating human pressures on protected areas rely mainly on land-use changes such as those used in compiling the human footprint index (Jones et al., 2018). This index, which quantifies conversion of land to agriculture, urbanization, and human infrastructure, significantly underestimates (or overlooks) impacts of IAS in regions perceived to have low human pressures (Hulme, 2018). For example, it ignores the potential for triggering ecosystem regime shifts (Gaertner et al., 2014). An important question is how effective protected areas will be in protecting native species and ecosystems from impacts caused by invasions under accelerating climate change (Baron et al., 2009). By investigating current and future potential distributions of 100 of the most invasive terrestrial, freshwater, and marine species in Europe, Gallardo et al. (2017) evaluated the combined threat posed by invasions and climate change. They found that the predicted richness of IAS was 11–18% lower inside than outside protected areas. They concluded that protected areas can provide strategic refugia for native species and recommended prioritizing actions to protect them from incursions of IAS spreading under climate change. III. WHY SHOULD WE CARE? THE IMPACTS OF BIOLOGICAL INVASIONS (1) Environmental impacts Given the many pressing environmental issues, one must ask how prominently biological invasions should feature in political and public agendas? The magnitude and variety of impacts of IAS provide an unambiguous reason for much more urgent attention to be given to invasions. Research on invasion impacts has developed rapidly over the past decade (Ricciardi et al., 2013; Anton et al., 2019), yielding improved understanding of mechanisms underlying these impacts and development of a sound theoretical basis and conceptual frameworks (Jeschke et al., 2014; Kumschick et al., 2015b, 2017). Such advances have paved the way for developing tools for quantitative impact assessment and for Fig 2. Increase in cumulative established alien species richness across six taxonomic groups in four regions of the world. Time series are based on the year of first record of those alien species that later became established in the given region (based on Seebens et al., 2017). Biological Reviews 95 (2020) 1511–1534 © 2020 The Authors. Biological Reviews published by John Wiley & Sons Ltd on behalf of Cambridge Philosophical Society. 1518 Petr Pyšek et al. the practical application of such protocols in biodiversity conservation and environmental management (Blackburn et al., 2014; Hawkins et al., 2015; Nentwig et al., 2016). Invasive alien species affect native species richness and abundance (Vilà et al., 2011; Pyšek et al., 2012; Kumschick et al., 2015a; Cameron, Vilà, & Cabeza, 2016; Gallardo et al., 2016) and have broken down biogeographical realms (Capinha et al., 2015), and they hinder ecosystem functioning and provision of ecosystem services (Gaertner et al., 2014; Vilà & Hulme, 2017; Castro-Díez et al., 2019). They can increase the risk of native species extinction, affect the genetic composition of native populations, modify the phylogenetic and functional diversity of invaded communities and trophic networks, and alter ecosystem productivity, nutrient and contaminant cycling, hydrology, and disturbance regimes (e.g. Brooks et al., 2004; Suarez & Tsutsui, 2008; Kenis et al., 2009; Ricciardi et al., 2013; Blackburn, Bellard, & Ricciardi, 2019). Impacts of alien species vary greatly across species, regions, and ecosystems (Blackburn et al.,2014)anddependontheabundance of the alien species and their trophic levels relative to those of affected native species (Hejda, Pyšek, & Jarošík, 2009; Bradley et al., 2019). For invasive plants, 63% of studies that have measuredimpactsfoundsignificant differences in species, community, or ecosystem characteristics compared to the situation prior to invasion, and impacts are far more likely to occur on resident plant and animal richness on islands than on mainlands (Pyšek et al., 2012). Many invasive alien plants modify ecosystems in ways that enhance their own persistence and suppress native species through reinforcing feedbacks, causing regime shifts (altered states of ecosystem structure and function that are difficult or impossible to reverse). Examples include impacts on soil-nutrient cycling caused by invasive trees and shrubs in forests and by herbaceous invaders in wetlands, through modifying the composition of soil seed banks and changed fire regimes (Gaertner et al., 2014; Gioria, Jarošík, & Pyšek, 2014; Shackleton et al., 2018) and altering microbial communities (Bowen et al., 2017). The IAS with the greatest impacts emerge from all taxonomic groups, as illustrated by the example of Europe. The European list of aliens with highest impacts includes 149 species: 54 plants, 49 invertebrates, 40 vertebrates, and six fungi. Among the highest-ranking species are one bird species (Canada goose Branta canadensis), four mammals (Norway rat Rattus norvegicus,muskratOndatra zibethicus, Sika deer Cervus nippon,Reeve’s muntjac Muntiacus reevesi), one crayfish (Procambarus clarkii), the varroa mite (Varroa destructor), and four plants (silver wattle Acacia dealbata,redsageLantana camara,kudzuPueraria lobata,waterhyacinth Eichhornia crassipes)(Nentwiget al., 2018). Extinctions owing to IAS constitute a special case of the ultimate threat to biodiversity and conservation (Bellard, Cassey, & Blackburn, 2016; Blackburn, Bellard, & Ricciardi, 2019). Invasive alien species are listed as one driver of extinction for 261 of 782 animal species and in 39 of 153 plant species worldwide; for both groups, IAS rank as the mostfrequentcause,aheadof hunting, harvesting, and agriculture (Blackburn, Bellard, & Ricciardi, 2019). The most vulnerable species are island endemics that have limited experience of mammalian predators or herbivoresandnowheretoescapeto[seePyšek et al., 2017a; Blackburn, Bellard, & Ricciardi, 2019 and references therein]. Observational evidence comparing alien plants, mammals, reptiles, fishes, molluscs, earthworms, and insects as causes of population declines or extinctions of native taxa suggests that alien predators are far more likely than alien competitors to cause the extinction of native species (Pyšek et al., 2017a). Notable predators include alien vertebrates and molluscs (Table 1). Plants (e.g. red cinchona Cinchona pubescens on the Galapágos Islands, strawberry guava Psidium cattleianum on Mauritius) and insects (e.g. cactus moth Cactoblastis cactorum in North America, harlequin ladybird Harmonia axyridis in Europe; Fig. 3) are all known to reduce population sizes of native species. Several fungal pathogens also significantly affect their host species in the new ranges (e.g. Batrachochytrium dendrobatidis on amphibians, Ophiostoma novo-ulmi on European elm trees Ulmus minor). One must also consider that native species, even if not yet driven to local or global extinction, often suffer from population declines attributable to IAS, and many now exist only as remnant populations (Downey & Richardson, 2016; Pyšek et al., 2017a). These declines can also cause interspecific interactions to be lost long before species disappear, affecting ecosystem function and services more severely than would be inferred from the rate of species extinctions (Valiente-Banuet et al., 2015). Thus, calls to downplay invasion impacts by citing short-term regional increases in total biodiversity caused by alien species are misleading, as are suggestions that losses caused by invasions will be counterbalanced by ‘new speciation’(Richardson & Ricciardi, 2013; Pauchard et al., 2018). An argument that appears occasionally in the literature is that the impacts of alien species on biodiversity and ecosystem functioning are similar to those of widespread, dominant native species [e.g. Davis et al., 2011, but see Simberloff et al., 2011]. However, contrary evidence is accumulating (e.g. Paolucci, MacIsaac, & Ricciardi, 2013; Buckley & Catford, 2016; Hejda, Štajerová, & Pyšek, 2017). A recent analysis of data on global extinctions in the IUCN Red List database (IUCN, 2017) revealed that alien species contributed to 25% of plant extinctions and 33% of terrestrial and freshwater animal extinctions; these figures are an order of magnitude higher than for native species, which were implicated in fewer than 5% and 3% of plant and animal extinctions, respectively (Blackburn, Bellard, & Ricciardi, 2019). For the USA, established alien plant species are 40 times more likely to be problematic for local ecosystems than are native species (Simberloff et al., 2012). The impact of invasions driving biodiversity change must be considered not only on its own (as are drivers listed by Ripple et al., 2017) but in concert with other drivers, such as climate or land-use change (Bradford et al., 2007; Walther et al., 2009; Schweiger et al., 2010; Hulme, 2011b; Chytrý et al., 2012). One example of an exacerbating interaction is the predicted future effect of climate, socioeconomic factors, and invasions on biodiversity hotspots. Socioeconomic factors, such as trade, have played a key role in the recent rapid spread of alien species. A study combining data on 60-year trends of bilateral trade among 147 countries with trends in biodiversity and climate showed particularly strong increases in established Biological Reviews 95 (2020) 1511–1534 © 2020 The Authors. Biological Reviews published by John Wiley & Sons Ltd on behalf of Cambridge Philosophical Society. Global overview of biological invasions 1519 IPBES Plenary composed of representatives from 132 member states. (2) Management interventions need to be objectively prioritized Invasions are pervasive –thousands of alien species have arrived, and more are arriving almost everywhere –and require much bolder actions. We can manage the most significant IAS and protect the most vulnerable ecosystems, but this requires a significant leap of commitment. It is important to prioritize, for example by focusing on vulnerable areas that are most at risk, including in developing countries, some of which are megadiversity hotspots (Seebens et al., 2015) or islands (Dawson et al., 2017). This paves the way for improved efficiency by focusing management on (i) high-risk pathways, activities, and/or societal sectors that use alien species (e.g. commercial forestry, ornamental horticulture, biofuels, pet trade, shipping) to prevent introduction, (ii) newly arrived species for removal to prevent further spread, and (iii) vulnerable habitats/native species to monitor and impede invasions from impacting them. Managing invasions is difficult and often expensive, but emerging evidence shows that even expensive interventions, especially related to prevention, generally result in net benefits (Zavaleta, 2000; Keller, Lodge, & Finnoff, 2007). (3) Protected areas need special attention Protected areas are an important part of global efforts to conserve biodiversity. Nevertheless, integrated efforts involving science, management, and policy for dealing with IAS in protected areas are insufficient. The extent and overall impact of invasive species in protected areas is increasing worldwide, especially for invasive plants, despite some notable successes in dealing with such invasions (Shackleton et al., 2020). New initiatives are needed to pave the way for monitoring trends, revising legislation and policies, and improving management interventions to reduce the impacts of invasive alien species in protected areas (Genovesi & Monaco, 2013; Foxcroft et al., 2017). However, management actions are also needed in human-dominated systems, including urban ecosystems where invasions are becoming increasingly problematic for human well-being and from which invasions into protected areas are often launched. (4) More effective protocols are needed for engaging with the public and societal actors Invasive alien species are affecting many aspects of human society, and their management requires the involvement of all societal stakeholders. Strengthening multidisciplinary approaches to invasion science (Simberloff et al., 2013; Vaz et al., 2017) is becoming a conditio sine qua non in the quest for comprehensive solutions to deal with biological invasions. Overcoming knowing–doing gaps (Hulme, 2014a; Foxcroft et al., 2020), involving citizen science (Groom et al., 2019), and raising awareness are making important contributions towards developing effective, operational, and clear mechanisms for much greater public and, hence, political engagement with the many complex and interacting dimensions of biological invasions. An under-developed area is engaging indigenous perspectives on the threat of alien species to culture and livelihoods and how to manage them. More efforts are needed to understand how IAS are directly affecting human well-being (i.e. Good Quality of Life in the IPBES framework) and how management can reduce such impacts. (5) Forecasting and scenario development must give more attention to synergies of invasions with climate change and other environmental changes Despite numerous correlative bioclimatic models predicting that many alien species will likely become more widespread as a result of climate change, there is a dearth of empirical data that clearly link shifts in alien species distributions with changes in temperature or precipitation. Analysis of longterm and large-scale spatial data on alien species distributions is urgently needed to disentangle how they correlate with climate variables and other aspects of global change, such as intensified land use and transformation, pollution, and propagule pressure (Bellard et al., 2013; González-Mor- eno et al., 2014; Mazor et al., 2018). Emerging research shows that we must address synergies in interacting drivers of invasions, synergies in impacts of multiple invaders, and species interactions resulting in invasional meltdowns and other feedbacks, as well as regime shifts (Gaertner et al., 2014), to improve our ability to predict new invasions and their impacts. VII. CONCLUSIONS (1) Biological invasions are a major driver of ecosystem degradation. The number of invasive alien species is increasing rapidly with no evidence that either the rate of species introduction or the emergence of new invasive species is slowing down. (2) Islands and coastal mainland areas are hotspots of invasions, but ecosystems in all biomes throughout the world are increasingly affected. Although boundaries of protected areas provide some resistance to invasions, even the most isolated and well-managed reserves are experiencing pressure from invasive alien species. (3) The global escalation in biological invasions is attributed to the increase in the number of pathways of introduction and spread of species, and particularly the volume of traffic (and therefore species) along these pathways. Emerging pathways are creating new categories of invasions, such as plastics providing rafts for transport of organisms across oceans. (4) Interactions with other drivers of global change are exacerbating current biological invasions and facilitating Biological Reviews 95 (2020) 1511–1534 © 2020 The Authors. Biological Reviews published by John Wiley & Sons Ltd on behalf of Cambridge Philosophical Society. 1526 Petr Pyšek et al. new ones, thereby greatly escalating the extent and impacts of invaders. Although biological invasions are sometimes symptoms (or ‘passengers’) of other human-mediated change, they are themselves often key drivers of change. (5) Invasions have complex and often immense long-term direct and indirect impacts, many of which manifest decades or more after invasions commence, when the invaders are established and extend across large geographic ranges. Invasive alien species break down biogeographic realms, affect native species richness and abundance, increase the risk of native species extinction, affect the genetic composition of native populations, change native animal behaviour, alter phylogenetic diversity across communities, and modify trophic networks. Many invasions alter ecosystem functioning and the delivery of ecosystem services, thereby adversely impacting human livelihoods. All these types of impacts are accelerating and predicted to increase in the future, often following non-linear trajectories. Despite advances in understanding impacts of biological invasions, little is known about the impacts of alien pathogens (including viruses, bacteria, fungi, and protists) and associated emerging infectious diseases on biodiversity and ecosystems. (6) Strategies to reduce future invasions are in place in many countries but are often implemented ineffectively. Unlike some other facets of global environmental change, with sufficient foresight and resources many biological invasions can be managed and mitigated. There is increasing evidence of successful long-term and large-scale management of established invaders, such as the eradication of mammals on increasingly large islands and biological control of weeds across continental areas. In many countries, however, invasions receive inadequate attention. Management approaches must be objectively prioritized by accounting for feasibility and considering invasion debt (the delayed spread of species after introduction to a region and the inevitable escalation of impacts over time). Engaging people from diverse stakeholder groups is essential to enhance understanding of biological invasions and inform decision-making including effective implementation of biosecurity. (7) Multidisciplinary collaborations and integrated approaches through international cooperation are critical to reduce the impacts of invasive alien species, including alien pathogens, on biodiversity, ecosystem services, and human livelihoods. Countries must strengthen their biosecurity regulations to implement and enforce effective management strategies that address invasions in tandem with other facets of global change. VIII. ACKNOWLEDGEMENTS P.P., J.P. and L.C.F. were supported by EXPRO grant no. 19-28807X (Czech Science Foundation) and long-term research development project RVO 67985939 (Czech Academy of Sciences). S.B. was supported through the 2017-2018 Belmont Forum and joint call for research proposals, under the BiodivScen ERA-Net COFUND programme, and by the Swiss National Science Foundation (grant numbers 31BD30_184114, and 31003A_179491). F.E. was supported by a grant from the Austrian Science Foundation FWF (grant I 3757-B29). L.C.F. acknowledges South African National Parks, the DSI-NRF Centre of Excellence for Invasion Biology, Stellenbosch University, and the National Research Foundation of South Africa (Grant Numbers IFR2010041400019 and IFR160215158271). P.E.H. was supported through grant C09X1611 “Winning against Wildings”from the New Zealand Ministry of Business, Innovation and Employment. J.M.J. was supported by the Deutsche Forschungsgemeinschaft (DFG; grant JE 288/9-2) and the Belmont Forum-BiodivERsA projects InvasiBES and AlienScenarios with the national funder German Federal Ministry of Education and Research (BMBF; grants 01LC1803A and 01LC1807B). A.M.L. was supported by EVA4.0, No. CZ.02.1.01/0.0/0.0/16_019/0000803 financed by OP RDE and by the USDA Forest Service. A. P. was funded by CONICYT AFB-170008 and Fondecyt 1180205. D.M.R. acknowledges support from the DSINRF Centre of Excellence for Invasion Biology and the Oppenheimer Memorial Trust (grant 18576/03). H.E.R. was supported by the Natural Environment Research Council award number NE/R016429/1 as part of the UKSCAPE programme Delivering National Capability. M.v. K. was supported by the Deutsche Forschungsgemeinschaft (DFG: grant 264740629). M.V. was supported by the Belmont Forum-BiodivERsA project InvasiBES (PCI2018- 092939) funded by the Spanish Ministerio de Ciencia, Innovación y Universidades. 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