Although research on human-mediated exchanges of species has substantially intensified during the last centuries, we know surprisingly little about temporal dynamics of alien species accumulations across regions and taxa. Using a novel database of 45,813 first records of 16,926 established alien species, we show that the annual rate of first records worldwide has increased during the last 200 years, with 37% of all first records reported most recently (1970–2014). Inter-continental and inter-taxonomic variation can be largely attributed to the diaspora of European settlers in the nineteenth century and to the acceleration in trade in the twentieth century. For all taxonomic groups, the increase in numbers of alien species does not show any sign of saturation and most taxa even show increases in the rate of first records over time. This highlights that past efforts to mitigate invasions have not been effective enough to keep up with increasing globalization.

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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.

What determines invasiveness of alien organisms is among the most interesting and urgent questions in ecology. In attempts to answer this question, researchers compare invasive alien species either to native species or to non-invasive alien species, and this is done in either the introduced or native ranges. However, inferences that can be drawn from these comparisons differ considerably, and failure to recognize this could hamper the search for determinants of invasiveness. To increase awareness about this issue, we present a framework of the various comparisons that can be used to test for determinants of invasiveness, and the specific questions each comparison can address. Moreover, we discuss how different comparisons complement each other, and therefore should be used in concert. For progress in invasion biology, it is crucial to realize that different comparisons address different biological questions and that some questions can only be answered unambiguously by combining them.

Human-mediated transport beyond biogeographic barriers has led to the introduction and 73The transport of species across biogeographic barriers by humans is a key component of 74 global environmental change [1][2][3] . Some of the species introduced to new regions will establish 75 self-sustaining populations and, thus, become a persistent part of the local biota 95We expect regions with higher gross domestic product per capita (GDPpc) or with higher 96 population densities to receive more alien species introductions across taxa (i.e., to experience 97 higher colonisation pressure through trade and transport), resulting in higher EAS richness 7,8,10,21 . 98We also test whether EAS richness patterns follow the latitudinal gradients often observed for 99 native biota, with higher richness in regions with higher mean annual temperature and 100 precipitation 22,23 . We expect island regions to have higher EAS richness than mainland regions, 101as islands are thought to be more prone to the establishment of alien species 12,24,25 . In addition, 102we expect more isolated oceanic islands to have greater EAS richness, as they have been shown 103 to receive more introductions, at least for birds 9 . We also expect coastal regions (as points of human population density, with a weak trend of higher alien richness in wetter regions (Table 1). 125While we only have potential proxy data (GDPpc, population density) for colonisation pressure 126 here (i.e., the total numbers of species introduced) 26 , our results suggest that cumulative numbers 127 7 of EAS are driven to a greater extent by differences in area and the pressure of introductions 128 from human history and activity 1,3,5,12,21 than by climate. 129Island regions have on average higher cross-taxon EAS richness (mean ± 1 S.D. 130proportional cross-taxon richness = 0.17 ± 0.11) than mainland regions (mean ± 1 S.D. = 0.11 ± 131 0.07; Table 1). In addition, models explaining alien richness of island and mainland regions 132 separately reveal that EAS richness is more strongly related to area, GDPpc and population 133 density on islands than in mainland regions (Table 1) (Table 1). Among mainland regions, EAS richness is greater for coastal (mean ± 1 S.D. 139proportional cross-taxon richness = 0.13 ± 0.09) than for landlocked regions (mean ± 1 S.D. = 140 0.10 ± 0.04). Cross-taxon EAS richness on islands tends to be higher for those further from 141 continental landmasses (Table 1). 143 Taxonomic congruence 144The strongest correlations in alien richness between taxonomic groups exist for ants and 145 reptiles (r s = 0.62), followed by birds and mammals, and vascular plants and spiders (both r s = 146 0.55) ( Table 2). For ants and reptiles, EAS richness is high in the Hawaiian Islands, southern 147United States (especially Florida) and Madagascar and the Mascarene Islands (Fig. 1b, 1g). (Fig. 1f, 1h). In Europe, the United Kingdom has the highest established alien 154 plant richness, while Germany has the highest spider richness (Fig. 1h, 1h). Overa...

Our ability to predict the identity of future invasive alien species is largely based upon knowledge of prior invasion history. Emerging alien species-those never encountered as aliens before-therefore pose a significant challenge to biosecurity interventions worldwide. Understanding their temporal trends, origins, and the drivers of their spread is pivotal to improving prevention and risk assessment tools. Here, we use a database of 45,984 first records of 16,019 established alien species to investigate the temporal dynamics of occurrences of emerging alien species worldwide. Even after many centuries of invasions the rate of emergence of new alien species is still high: One-quarter of first records during 2000-2005 were of species that had not been previously recorded anywhere as alien, though with large variation across taxa. Model results show that the high proportion of emerging alien species cannot be solely explained by increases in well-known drivers such as the amount of imported commodities from historically important source regions. Instead, these dynamics reflect the incorporation of new regions into the pool of potential alien species, likely as a consequence of expanding trade networks and environmental change. This process compensates for the depletion of the historically important source species pool through successive invasions. We estimate that 1-16% of all species on Earth, depending on the taxonomic group, qualify as potential alien species. These results suggest that there remains a high proportion of emerging alien species we have yet to encounter, with future impacts that are difficult to predict.

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Biological invasions have steadily increased over recent centuries. However, we still lack a clear expectation about future trends in alien species numbers. In particular, we do not know whether alien species will continue to accumulate in regional floras and faunas, or whether the pace of accumulation will decrease due to the depletion of native source pools. Here, we apply a new model to simulate future numbers of alien species based on estimated sizes of source pools and dynamics of historical invasions, assuming a continuation of processes in the future as observed in the past (a business‐as‐usual scenario). We first validated performance of different model versions by conducting a back‐casting approach, therefore fitting the model to alien species numbers until 1950 and validating predictions on trends from 1950 to 2005. In a second step, we selected the best performing model that provided the most robust predictions to project trajectories of alien species numbers until 2050. Altogether, this resulted in 3,790 stochastic simulation runs for 38 taxon–continent combinations. We provide the first quantitative projections of future trajectories of alien species numbers for seven major taxonomic groups in eight continents, accounting for variation in sampling intensity and uncertainty in projections. Overall, established alien species numbers per continent were predicted to increase from 2005 to 2050 by 36%. Particularly, strong increases were projected for Europe in absolute (+2,543 ± 237 alien species) and relative terms, followed by Temperate Asia (+1,597 ± 197), Northern America (1,484 ± 74) and Southern America (1,391 ± 258). Among individual taxonomic groups, especially strong increases were projected for invertebrates globally. Declining (but still positive) rates were projected only for Australasia. Our projections provide a first baseline for the assessment of future developments of biological invasions, which will help to inform policies to contain the spread of alien species.