Regional invasion history and land use shape the prevalence of non-native species in local assemblages.

The ecological impact of non-native species arises from their establishment in local assemblages. However, the rates of non-native spread in new regions and their determinants have not been comprehensively studied. Here, we combined global databases documenting the occurrence of non-native species and residence of non-native birds, mammals, and vascular plants at regional and local scales to describe how the likelihood of non-native occurrence and their proportion in local assemblages relate with their residence time and levels of human usage in different ecosystems. Our findings reveal that local non-native occurrence generally increases with residence time. Colonization is most rapid in croplands and urban areas, while it is slower and variable in natural or semi-natural ecosystems. Notably, non-native occurrence continues to rise even 200 years after introduction, especially for birds and vascular plants, and in other land-use types rather than croplands and urban areas. The impact of residence time on non-native proportions is significant only for mammals. We conclude that the continental exchange of biotas requires considerable time for effects to manifest at the local scale across taxa and land-use types. The unpredictability of future impacts, implied by the slow spread of non-native species, strengthens the call for stronger regulations on the exchange of non-native species to reduce the long-lasting invasion debt looming on ecosystems' future.

Long-term trajectories of non-native vegetation on islands globally.

Human-mediated changes in island vegetation are, among others, largely caused by the introduction and establishment of non-native species. However, data on past changes in non-native plant species abundance that predate historical documentation and censuses are scarce. Islands are among the few places where we can track human arrival in natural systems allowing us to reveal changes in vegetation dynamics with the arrival of non-native species. We matched fossil pollen data with botanical status information (native, non-native), and quantified the timing, trajectories and magnitude of non-native plant vegetational change on 29 islands over the past 5000 years. We recorded a proportional increase in pollen of non-native plant taxa within the last 1000 years. Individual island trajectories are context-dependent and linked to island settlement histories. Our data show that non-native plant introductions have a longer and more dynamic history than is generally recognized, with critical implications for biodiversity baselines and invasion biology.

Small genome size and variation in ploidy levels support the naturalization of vascular plants but constrain their invasive spread.

Karyological characteristics are among the traits underpinning the invasion success of vascular plants. Using 11 049 species, we tested the effects of genome size and ploidy levels on plant naturalization (species forming self-sustaining populations where they are not native) and invasion (naturalized species spreading rapidly and having environmental impact). The probability that a species naturalized anywhere in the world decreased with increasing monoploid genome size (DNA content of a single chromosome set). Naturalized or invasive species with intermediate monoploid genomes were reported from many regions, but those with either small or large genomes occurred in fewer regions. By contrast, large holoploid genome sizes (DNA content of the unreplicated gametic nucleus) constrained naturalization but favoured invasion. We suggest that a small genome is an advantage during naturalization, being linked to traits favouring adaptation to local conditions, but for invasive spread, traits associated with a large holoploid genome, where the impact of polyploidy may act, facilitate long-distance dispersal and competition with other species.

Patterns and drivers of climatic niche dynamics during biological invasions of island-endemic amphibians, reptiles, and birds.

Shifts between native and alien climatic niches pose a major challenge for predicting biological invasions. This is particularly true for insular species because geophysical barriers could constrain the realization of their fundamental niches, which may lead to underestimates of their invasion potential. To investigate this idea, we estimated the frequency of shifts between native and alien climatic niches and the magnitude of climatic mismatches using 80,148 alien occurrences of 46 endemic insular amphibian, reptile, and bird species. Then, we assessed the influence of nine potential predictors on climatic mismatches across taxa, based on species' characteristics, native range physical characteristics, and alien range properties. We found that climatic mismatch is common during invasions of endemic insular birds and reptiles: 78.3% and 55.1% of their respective alien records occurred outside of the environmental space of species' native climatic niche. In comparison, climatic mismatch was evident for only 16.2% of the amphibian invasions analyzed. Several predictors significantly explained climatic mismatch, and these varied among taxonomic groups. For amphibians, only native range size was associated with climatic mismatch. For reptiles, the magnitude of climatic mismatch was higher for species with narrow native altitudinal ranges, occurring in topographically complex or less remote islands, as well as for species with larger distances between their native and alien ranges. For birds, climatic mismatch was significantly larger for invasions on continents with higher phylogenetic diversity of the recipient community, and when the invader was more evolutionarily distinct. Our findings highlight that apparently common niche shifts of insular species may jeopardize our ability to forecast their potential invasions using correlative methods based on climatic variables. Also, we show which factors provide additional insights on the actual invasion potential of insular endemic amphibians, reptiles, and birds.

Human appropriation of net primary production as driver of change in landscape-scale vertebrate richness.

Aim: Land use is the most pervasive driver of biodiversity loss. Predicting its impact on species richness (SR) is often based on indicators of habitat loss. However, the degradation of habitats, especially through land-use intensification, also affects species. Here, we evaluate whether an integrative metric of land-use intensity, the human appropriation of net primary production, is correlated with the decline of SR in used landscapes across the globe. Location: Global. Time period: Present. Major taxa studied: Birds, mammals and amphibians. Methods: Based on species range maps (spatial resolution: 20 km × 20 km) and an area-of-habitat approach, we calibrated a "species-energy model" by correlating the SR of three groups of vertebrates with net primary production and biogeographical covariables in "wilderness" areas (i.e., those where available energy is assumed to be still at pristine levels). We used this model to project the difference between pristine SR and the SR corresponding to the energy remaining in used landscapes (i.e., SR loss expected owing to human energy extraction outside wilderness areas). We validated the projected species loss by comparison with the realized and impending loss reconstructed from habitat conversion and documented by national Red Lists. Results: Species-energy models largely explained landscape-scale variation of mapped SR in wilderness areas (adjusted R 2-values: 0.79-0.93). Model-based projections of SR loss were lower, on average, than reconstructed and documented ones, but the spatial patterns were correlated significantly, with stronger correlation in mammals (Pearson's r = 0.68) than in amphibians (r = 0.60) and birds (r = 0.57). Main conclusions: Our results suggest that the human appropriation of net primary production is a useful indicator of heterotrophic species loss in used landscapes, hence we recommend its inclusion in models based on species-area relationships to improve predictions of land-use-driven biodiversity loss.

Economic costs of protecting islands from invasive alien species.

Biological invasions represent a key threat to insular systems and have pronounced impacts across environments and economies. The ecological impacts have received substantial focus, but the socioeconomic impacts are poorly synthesized across spatial and temporal scales. We used the InvaCost database, the most comprehensive assessment of published economic costs of invasive species, to assess economic impacts on islands worldwide. We analyzed socioeconomic costs across differing expenditure types and examined temporal trends across islands that differ in their political geography-island nation states, overseas territories, and islands of continental countries. Over US$36 billion in total costs (including damages and management) has occurred on islands from 1965 to 2020 due to invasive species' impacts. Nation states incurred the greatest total and management costs, and islands of continental countries incurred costs of similar magnitude, both far higher than those in overseas territories. Damage-loss costs were significantly lower, but with qualitatively similar patterns across differing political geographies. The predominance of management spending differs from the pattern found for most countries examined and suggests important knowledge gaps in the extent of many damage-related socioeconomic impacts. Nation states spent the greatest proportion of their gross domestic products countering these costs, at least 1 order of magnitude higher than other locations. Most costs were borne by authorities and stakeholders, demonstrating the key role of governmental and nongovernmental bodies in addressing island invasions. Temporal trends revealed cost increases across all island types, potentially reflecting efforts to tackle invasive species at larger, more socially complex scales. Nevertheless, the already high total economic costs of island invasions substantiate the role of biosecurity in reducing and preventing invasive species arrivals to reduce strains on limited financial resources and avoid threats to sustainable development goals.

Costos económicos de proteger a las islas de las especies invasoras Resumen Las invasiones biológicas representan una amenaza importante para los sistemas insulares, además de tener impactos pronunciados en el ambiente y en la economía. Los impactos ecológicos han recibido atención sustancial, mientras que los impactos socioeconómicos se encuentran pobremente sintetizados en las escalas temporales y espaciales. Usamos la base de datos InvaCost, el análisis más completo de los costos económicos de las especies invasoras, para evaluar los impactos económicos sobre las islas a nivel mundial. Analizamos los costos socioeconómicos en varios tipos de gastos y examinamos las tendencias temporales en las islas que difieren en su geografía política - islas estado-nación, territorios ultramarinos e islas de países continentales. En las islas han ocurrido gastos de más de $36 mil millones de dólares entre 1965 y 2020 debido a los impactos de las especies invasoras. Las islas estado-nación produjeron los mayores costos de manejo y el mayor total, mientras que las islas de los países continentales produjeron costos de una magnitud similar, ambas con gastos mucho más elevados que los de los territorios ultramarinos. Los costos de las pérdidas por daños fueron significativamente más bajas, aunque con patrones cualitativamente similares entre las diferentes geografías políticas. El predominio del gasto en el manejo difiere del patrón hallado en la mayoría de los países analizados y sugiere que hay vacíos importantes en el conocimiento del alcance de muchos de los impactos socioeconómicos relacionados con los daños. Las islas estado-nación gastaron la mayor proporción de su producto interno bruto en contrarrestar estos costos, al menos una orden de magnitud mayor que las otras localidades. La mayoría de los costos fueron asumidos por las autoridades y los accionistas, lo que demuestra el papel clave que tienen los organismos gubernamentales y no gubernamentales en cómo se atienden las invasiones insulares. Las tendencias temporales revelaron incrementos en el costo en todos los tipos de islas, lo que potencialmente refleja los esfuerzos por combatir a las especies invasoras a escalas más grandes y socialmente más complejas. Aun así, el elevado costo económico total de las invasiones insulares fundamenta la función que tiene la bioseguridad en la reducción y prevención de la llegada de especies invasoras para reducir presiones sobre los recursos financieros limitados y evitar amenazas para las metas de desarrollo sustentable.

Role of diversification rates and evolutionary history as a driver of plant naturalization success.

Human introductions of species beyond their natural ranges and their subsequent establishment are defining features of global environmental change. However, naturalized plants are not uniformly distributed across phylogenetic lineages, with some families contributing disproportionately more to the global alien species pool than others. Additionally, lineages differ in diversification rates, and high diversification rates have been associated with characteristics that increase species naturalization success. Here, we investigate the role of diversification rates in explaining the naturalization success of angiosperm plant families. We use five global data sets that include native and alien plant species distribution, horticultural use of plants, and a time-calibrated angiosperm phylogeny. Using phylogenetic generalized linear mixed models, we analysed the effect of diversification rate, different geographical range measures, and horticultural use on the naturalization success of plant families. We show that a family's naturalization success is positively associated with its evolutionary history, native range size, and economic use. Investigating interactive effects of these predictors shows that native range size and geographic distribution additionally affect naturalization success. High diversification rates and large ranges increase naturalization success, especially of temperate families. We suggest this may result from lower ecological specialization in temperate families with large ranges, compared with tropical families with smaller ranges.

Remoteness promotes biological invasions on islands worldwide.

One of the best-known general patterns in island biogeography is the species-isolation relationship (SIR), a decrease in the number of native species with increasing island isolation that is linked to lower rates of natural dispersal and colonization on remote oceanic islands. However, during recent centuries, the anthropogenic introduction of alien species has increasingly gained importance and altered the composition and richness of island species pools. We analyzed a large dataset for alien and native plants, ants, reptiles, mammals, and birds on 257 (sub) tropical islands, and showed that, except for birds, the number of naturalized alien species increases with isolation for all taxa, a pattern that is opposite to the negative SIR of native species. We argue that the reversal of the SIR for alien species is driven by an increase in island invasibility due to reduced diversity and increased ecological naiveté of native biota on the more remote islands.

Global rise in emerging alien species results from increased accessibility of new source pools.

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.

Projecting the continental accumulation of alien species through to 2050.

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.

Drivers of future alien species impacts: An expert-based assessment.

Understanding the likely future impacts of biological invasions is crucial yet highly challenging given the multiple relevant environmental, socio-economic and societal contexts and drivers. In the absence of quantitative models, methods based on expert knowledge are the best option for assessing future invasion trajectories. Here, we present an expert assessment of the drivers of potential alien species impacts under contrasting scenarios and socioecological contexts through the mid-21st century. Based on responses from 36 experts in biological invasions, moderate (20%-30%) increases in invasions, compared to the current conditions, are expected to cause major impacts on biodiversity in most socioecological contexts. Three main drivers of biological invasions-transport, climate change and socio-economic change-were predicted to significantly affect future impacts of alien species on biodiversity even under a best-case scenario. Other drivers (e.g. human demography and migration in tropical and subtropical regions) were also of high importance in specific global contexts (e.g. for individual taxonomic groups or biomes). We show that some best-case scenarios can substantially reduce potential future impacts of biological invasions. However, rapid and comprehensive actions are necessary to use this potential and achieve the goals of the Post-2020 Framework of the Convention on Biological Diversity.

Invasion costs, impacts, and human agency: response to Sagoff 2020.

Article impact statement: In an era of profound biodiversity crisis, invasion costs, invader impacts, and human agency should not be dismissed.

The Global Naturalized Alien Flora (GloNAF) database.

This dataset provides the Global Naturalized Alien Flora (GloNAF) database, version 1.2. GloNAF represents a data compendium on the occurrence and identity of naturalized alien vascular plant taxa across geographic regions (e.g. countries, states, provinces, districts, islands) around the globe. The dataset includes 13,939 taxa and covers 1,029 regions (including 381 islands). The dataset is based on 210 data sources. For each taxon-by-region combination, we provide information on whether the taxon is considered to be naturalized in the specific region (i.e. has established self-sustaining populations in the wild). Non-native taxa are marked as "alien", when it is not clear whether they are naturalized. To facilitate alignment with other plant databases, we provide for each taxon the name as given in the original data source and the standardized taxon and family names used by The Plant List Version 1.1 (http://www.theplantlist.org/). We provide an ESRI shapefile including polygons for each region and information on whether it is an island or a mainland region, the country and the Taxonomic Databases Working Group (TDWG) regions it is part of (TDWG levels 1-4). We also provide several variables that can be used to filter the data according to quality and completeness of alien taxon lists, which vary among the combinations of regions and data sources. A previous version of the GloNAF dataset (version 1.1) has already been used in several studies on, for example, historical spatial flows of taxa between continents and geographical patterns and determinants of naturalization across different taxonomic groups. We intend the updated and expanded GloNAF version presented here to be a global resource useful for studying plant invasions and changes in biodiversity from regional to global scales. We release these data into the public domain under a Creative Commons Zero license waiver (https://creativecommons.org/share-your-work/public-domain/cc0/). When you use the data in your publication, we request that you cite this data paper. If GloNAF is a major part of the data analyzed in your study, you should consider inviting the GloNAF core team (see Metadata S1: Originators in the Overall project description) as collaborators. If you plan to use the GloNAF dataset, we encourage you to contact the GloNAF core team to check whether there have been recent updates of the dataset, and whether similar analyses are already ongoing.

A Framework for Global Twenty-First Century Scenarios and Models of Biological Invasions.

Biological invasions have emerged as an eminent feature of global change, with substantial impacts on the environment and human livelihoods. Current research demonstrates that the numbers and impacts of alien species are rising unabatedly. At the same time, we lack a thorough understanding of potential future trajectories for the decades to come. With the recent establishment of comprehensive global databases, it is, for the first time, feasible to develop and quantify future scenarios of biological invasions. Therefore, we propose a conceptual framework for how to develop alien species scenarios for the twenty-first century and how to identify relevant steps and challenges along the way. The concept will be important to inform research, policy, stakeholders, and the general public. Furthermore, we call for the scientific community to join forces and to operationalize the framework for scenarios and models of biological invasions to develop an important baseline for understanding and managing future biological invasions.

The role of adaptive strategies in plant naturalization.

Determining the factors associated with the naturalization of alien species is a central theme in ecology. Here, we tested the usefulness of a metric for quantifying Grime's seminal concept of adaptive strategies - competitors, stress-tolerators and ruderals (CSR) - to explain plant naturalizations worldwide. Using a global dataset of 3004 vascular plant species, and accounting for phylogenetic relatedness and species' native biomes, we assessed the associations between calculated C-, S- and R-scores and naturalization success for species exhibiting different life forms. Across different plant life forms, C-scores were positively and S-scores negatively associated with both the probability of naturalization and the number of regions where the species has naturalized. R-scores had positive effects on the probability of naturalization. These effects of the scores were, however, weak to absent for tree species. Our findings demonstrate the utility of CSR-score calculation to broadly represent, and potentially explain, the naturalization success of plant species.

Plant invasion and naturalization are influenced by genome size, ecology and economic use globally.

Human factors and plant characteristics are important drivers of plant invasions, which threaten ecosystem integrity, biodiversity and human well-being. However, while previous studies often examined a limited number of factors or focused on a specific invasion stage (e.g., naturalization) for specific regions, a multi-factor and multi-stage analysis at the global scale is lacking. Here, we employ a multi-level framework to investigate the interplay between plant characteristics (genome size, Grime's adaptive CSR-strategies and native range size) and economic use and how these factors collectively affect plant naturalization and invasion success worldwide. While our findings derived from structural equation models highlight the substantial contribution of human assistance in both the naturalization and spread of invasive plants, we also uncovered the pivotal role of species' adaptive strategies among the factors studied, and the significantly varying influence of these factors across invasion stages. We further revealed that the effects of genome size on plant invasions were partially mediated by species adaptive strategies and native range size. Our study provides insights into the complex and dynamic process of plant invasions and identifies its key drivers worldwide.

Global indicators of the environmental impacts of invasive alien species and their information adequacy.

Monitoring the extent to which invasive alien species (IAS) negatively impact the environment is crucial for understanding and mitigating biological invasions. Indeed, such information is vital for achieving Target 6 of the Kunming-Montreal Global Biodiversity Framework. However, to-date indicators for tracking the environmental impacts of IAS have been either lacking or insufficient. Capitalizing on advances in data availability and impact assessment protocols, we developed environmental impact indicators to track realized and potential impacts of IAS. We also developed an information status indicator to assess the adequacy of the data underlying the impact indicators. We used data on 75 naturalized amphibians from 82 countries to demonstrate the indicators at a global scale. The information status indicator shows variation in the reliability of the data and highlights areas where absence of impact should be interpreted with caution. Impact indicators show that growth in potential impacts are dominated by predatory species, while potential impacts from both predation and disease transmission are distributed worldwide. Using open access data, the indicators are reproducible and adaptable across scales and taxa and can be used to assess global trends and distributions of IAS, assisting authorities in prioritizing control efforts and identifying areas at risk of future invasions. This article is part of the theme issue 'Ecological novelty and planetary stewardship: biodiversity dynamics in a transforming biosphere'.

Potential sources of time lags in calibrating species distribution models.

The Anthropocene is characterized by a rapid pace of environmental change and is causing a multitude of biotic responses, including those that affect the spatial distribution of species. Lagged responses are frequent and species distributions and assemblages are consequently pushed into a disequilibrium state. How the characteristics of environmental change-for example, gradual 'press' disturbances such as rising temperatures due to climate change versus infrequent 'pulse' disturbances such as extreme events-affect the magnitude of responses and the relaxation times of biota has been insufficiently explored. It is also not well understood how widely used approaches to assess or project the responses of species to changing environmental conditions can deal with time lags. It, therefore, remains unclear to what extent time lags in species distributions are accounted for in biodiversity assessments, scenarios and models; this has ramifications for policymaking and conservation science alike. This perspective piece reflects on lagged species responses to environmental change and discusses the potential consequences for species distribution models (SDMs), the tools of choice in biodiversity modelling. We suggest ways to better account for time lags in calibrating these models and to reduce their leverage effects in projections for improved biodiversity science and policy.

Economic costs of invasive non-native species in urban areas: An underexplored financial drain.

Urbanization is an important driver of global change associated with a set of environmental modifications that affect the introduction and distribution of invasive non-native species (species with populations transported by humans beyond their natural biogeographic range that established and are spreading in their introduced range; hereafter, invasive species). These species are recognized as a cause of large ecological and economic losses. Nevertheless, the economic impacts of these species in urban areas are still poorly understood. Here we present a synthesis of the reported economic costs of invasive species in urban areas using the global InvaCost database, and demonstrate that costs are likely underestimated. Sixty-one invasive species have been reported to cause a cumulative cost of US$ 326.7 billion in urban areas between 1965 and 2021 globally (average annual cost of US$ 5.7 billion). Class Insecta was responsible for >99 % of reported costs (US$ 324.4 billion), followed by Aves (US$ 1.4 billion), and Magnoliopsida (US$ 494 million). The reported costs were highly uneven with the sum of the five costliest species representing 80 % of reported costs. Most reported costs were a result of damage (77.3 %), principally impacting public and social welfare (77.9 %) and authorities-stakeholders (20.7 %), and were almost entirely in terrestrial environments (99.9 %). We found costs reported for 24 countries. Yet, there are 73 additional countries with no reported costs, but with occurrences of invasive species that have reported costs in other countries. Although covering a relatively small area of the Earth's surface, urban areas represent about 15 % of the total reported costs attributed to invasive species. These results highlight the conservative nature of the estimates and impacts, revealing important biases present in the evaluation and publication of reported data on costs. We emphasize the urgent need for more focused assessments of invasive species' economic impacts in urban areas.

The poleward naturalization of intracontinental alien plants.

Plant introductions outside their native ranges by humans have led to substantial ecological consequences. While we have gained considerable knowledge about intercontinental introductions, the distribution and determinants of intracontinental aliens remain poorly understood. Here, we studied naturalized (i.e., self-sustaining) intracontinental aliens using native and alien floras of 243 mainland regions in North America, South America, Europe, and Australia. We revealed that 4510 plant species had intracontinental origins, accounting for 3.9% of all plant species and 56.7% of all naturalized species in these continents. In North America and Europe, the numbers of intracontinental aliens peaked at mid-latitudes, while the proportion peaked at high latitudes in Europe. Notably, we found predominant poleward naturalization, primarily due to larger native species pools in low-latitudes. Geographic and climatic distances constrained the naturalization of intracontinental aliens in Australia, Europe, and North America, but not in South America. These findings suggest that poleward naturalizations will accelerate, as high latitudes become suitable for more plant species due to climate change.