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Invasion biology examines species originated elsewhere and moved with the help of humans, and those species' impacts on biodiversity, ecosystem services, and human well-being. In a globalized world, the emergence and spread of many human infectious pathogens are quintessential biological invasion events. Some macroscopic invasive species themselves contribute to the emergence and transmission of human infectious agents. We review conceptual parallels and differences between human epidemics and biological invasions by animals and plants. Fundamental concepts in invasion biology regarding the interplay of propagule pressure, species traits, biotic interactions, eco-evolutionary experience, and ecosystem disturbances can help to explain transitions between stages of epidemic spread. As a result, many forecasting and management tools used to address epidemics could be applied to biological invasions and vice versa. Therefore, we advocate for increasing cross-fertilization between the two disciplines to improve prediction, prevention, treatment, and mitigation of invasive species and infectious disease outbreaks, including pandemics. Ministerio de Ciencia e Innovación PCI2018-092986, PCI2018-092939 Federal Ministry of Education and Research 01LC1803A Austrian Science Fund I 4011-B32 Comisión Nacional de Investigación Científica y Tecnológica AFB170008 Natural Environment Research Council NE/P016766/1 Gobierno de España RYC-2013-13445, RYC2018-025160-1 Natural Sciences and Engineering Research Council of Canada RGPIN-2016-03918

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: Onequarter 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. © 2018 National Academy of Sciences. All Rights Reserved.

Terrestrial, freshwater, and marine ecosystems are connected via multiple biophysical and ecological processes. Identifying and quantifying links among ecosystems is necessary for the uptake of integrated conservation actions across realms. Such actions are particularly important for species using habitats in more than one realm during their daily or life cycle. We reviewed information on the habitats of 2,408 species of European conservation concern and found that 30% of the species use habitats in multiple realms. Transportation and service corridors, which fragment species habitats, were identified as the most important threat impacting similar to 70% of the species. We examined information on 1,567 European Union (EU) conservation projects funded over the past 25 years, to assess the adequacy of efforts toward the conservation of "multi-realm" species at a continental scale. We discovered that less than a third of multi-realm species benefited from projects that included conservation actions across multiple realms. To achieve the EU's conservation target of halting biodiversity loss by 2020 and effectively protect multi-realm species, integrated conservation efforts across realms should be reinforced by: (1) recognizing the need for integrated management at a policy level, (2) revising conservation funding priorities across realms, and (3) implementing integrated land-freshwater-sea conservation planning and management.

Based on the ‘European Mires Book’ of the International Mire Conservation Group (IMCG), this article provides a composite map of national datasets as the first comprehensive peatland map for the whole of Europe. We also present estimates of the extent of peatlands and mires in each European country individually and for the entire continent. A minimum peat thickness criterion has not been strictly applied, to allow for (often historically determined) country-specific definitions. Our ‘peatland’ concept includes all ‘mires’, which are peatlands where peat is being formed. The map was constructed by merging national datasets in GIS while maintaining the mapping scales of the original input data. This ‘bottom-up’ approach indicates that the overall area of peatland in Europe is 593,727 km². Mires were found to cover more than 320,000 km² (around 54 % of the total peatland area). If shallow-peat lands (< 30 cm peat) in European Russia are also taken into account, the total peatland area in Europe is more than 1,000,000 km2 , which is almost 10 % of the total surface area. Composite inventories of national peatland information, as presented here for Europe, may serve to identify gaps and priority areas for field survey, and help to cross-check and calibrate remote sensing based mapping approaches. SI

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. © The Author(s) 2017.

Biological invasions can be associated with shifts of the species' climatic niches but the incidence of such shifts is under debate. The reproductive system might be a key factor controlling such shifts because it influences a species' evolutionary flexibility. However, the link between reproductive systems and niche dynamics in plant invasions has been little studied so far. We compiled global occurrence data sets of 13 congeneric sexual and apomictic species pairs, and used principal components analysis (PCA) and kernel smoothers to compare changes in climatic niche optima, breadths and unfilling/expansion between native and alien ranges. Niche change metrics were compared between sexual and apomictic species. All 26 species showed changes in niche optima and/or breadth and 14 species significantly expanded their climatic niches. However, we found no effect of the reproductive system on niche dynamics. Instead, species with narrower native niches showed higher rates of niche expansion in the alien ranges. Our results suggest that niche shifts are frequent in plant invasions but evolutionary potential may not be of major importance for such shifts. Niche dynamics rather appear to be driven by changes of the realized niche without adaptive change of the fundamental climatic niche. peerReviewed

Common ragweed (Ambrosia artemisiifolia) is an invasive weed native to North America producing very allergenic pollen which causes serious health effects like rhinitis, asthma and atopic dermatitis. It was introduced in Europe since the mid-19th century and invaded large areas during the last few decades (Pannonian plain, Northern Italy and South-Eastern France). Furthermore, there is a high potential for ragweed spread in current suitable habitats and future changes in Climate and land use may increase the spread by altering the climatic niche determined by physiological thresholds or affecting cropping patterns. The rate of spread depends also on seed dispersal due to natural or anthropogenic processes and the efficiency of ragweed eradication policies. However, ragweed airborne pollen concentrations depend not only on plant infestation, but also on phenology, pollen production, release, dispersion and atmospheric transport. Here, we present the first integrated modelling framework, based on an explicit representation of plant phenology, pollen production, and release to the atmosphere, to assess future changes in airborne pollen concentration under scenarios of climate and land use changes and seed dispersal. Two model suites are implemented differing in the atmospheric processing and in the driving climate models. The CHIMERE suite uses the Chemistry- Transport Model CHIMERE model, forced by regional climate simulations from the WRF model downscaling of the IPSL-CM5A-MR model. The RegCM suite uses the RegCM4 regional climate model forced by global climate simulations from HadGEM CMIP5. We performed three types of simulations (50 km grid covering Europe), which are hind-cast (2000–2012), historical (1986–2005) and future (2041–2060) simulations. The hindcast simulations, forced by ERAInterim reanalysis, are performed to calibrate and evaluate the modelling chain. The historical simulations are carried out using calibrated ragweed density to serve as a reference simulation for the future. We considered two contrasting RCPs (Representative Concentration Pathways) climate change scenarios including a high-end (RCP 8.5) and moderate (RCP 4.5) climate change scenarios and three seed dispersal scenarios (reference, slow and rapid). We show that airborne pollen concentrations may drastically increase in 2050 by a factor of 4.5 under highend (RCP 8.5) and 4.0 under moderate (RCP 4.5) climate change scenarios. This upsurge is largely dependent on the seed dispersal rate, making this increase vary in a range of factors from 2 to 12 according to the range of formulated assumptions. We estimate that about one third of the projected increases of pollen concentration are due to the on-going seed dispersal within the present niche regardless of climate change. Climate change will be responsible of two thirds of the future pollen loads increase. It will extend the habitat suitability for ragweed in Northern and Eastern Europe and result in higher pollen concentrations in established ragweed areas mostly due to a larger primary production with increasing CO2. Therefore, future increase of airborne pollen concentrations will be caused by the combined effects of climate change and ragweed seed dispersal in current and future suitable areas. Our results indicate that controlling the current European ragweed invasion will become more difficult in the future as the environment will be more favourable for ragweed growth and spread, highlighting the need for the development of effective and regionally co-ordinated eradication programmes.

n order to support the prioritisation of invasive alien species for future risk assessments, a horizon scanning methodology for the Europe was developed and implemented. The outcome was a list of 95 species, including all taxa (except microorganisms) within marine, terrestrial and freshwater environments, considered as very high or high priority for risk assessment.

The accelerating rates of international trade, travel, and transport in the latter half of the twentieth century have led to the progressive mixing of biota from across the world and the number of species introduced to new regions continues to increase. The importance of biogeographic, climatic, economic, and demographic factors as drivers of this trend is increasingly being realized but as yet there is no consensus regarding their relative importance. Whereas little may be done to mitigate the effects of geography and climate on invasions, a wider range of options may exist to moderate the impacts of economic and demographic drivers. Here we use the most recent data available from Europe to partition between macroecological, economic, and demographic variables the variation in alien species richness of bryophytes, fungi, vascular plants, terrestrial insects, aquatic invertebrates, fish, amphibians, reptiles, birds, and mammals. Only national wealth and human population density were statistically significant predictors in the majority of models when analyzed jointly with climate, geography, and land cover. The economic and demographic variables reflect the intensity of human activities and integrate the effect of factors that directly determine the outcome of invasion such as propagule pressure, pathways of introduction, eutrophication, and the intensity of anthropogenic disturbance. The strong influence of economic and demographic variables on the levels of invasion by alien species demonstrates that future solutions to the problem of biological invasions at a national scale lie in mitigating the negative environmental consequences of human activities that generate wealth and by promoting more sustainable population growth.

Recent comprehensive data provided through the DAISIE project (www.europe-aliens.org) have facilitated the development of the first pan-European assessment of the impacts of alien plants, vertebrates, and invertebrates in terrestrial, freshwater, and marine environments - on ecosystem services. There are 1094 species with documented ecological impacts and 1347 with economic impacts. The two taxonomic groups with the most species causing impacts are terrestrial invertebrates and terrestrial plants. The North Sea is the maritime region that suffers the most impacts. Across taxa and regions, ecological and economic impacts are highly correlated. Terrestrial invertebrates create greater economic impacts than ecological impacts, while the reverse is true for terrestrial plants. Alien species from all taxonomie groups affect "supporting", "provisioning", "regulating", and "cultural" services and interfere with human well-being. Terrestrial vertebrates are responsible for the greatest range of impacts, and these are widely distributed across Europe. Here, we present a review of the financial costs, as the first step toward calculating an estimate of the economic consequences of alien species in Europe. © The Ecological Society of America.

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—there-fore 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 dy-namics 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 any-where 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 impor-tant source regions. Instead, these dynamics reflect the incorpora-tion 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 histor-ically 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.

Human-mediated transport beyond biogeographic barriers has led to the introduction and establishment of alien species in new regions worldwide. However, we lack a global picture of established alien species richness for multiple taxonomic groups. Here, we assess global patterns and potential drivers of established alien species richness across eight taxonomic groups (amphib-ians, ants, birds, freshwater fishes, mammals, vascular plants, reptiles and spiders) for 186 islands and 423 mainland regions. Hotspots of established alien species richness are predominantly island and coastal mainland regions. Regions with greater gross domestic product per capita, human population density, and area have higher established alien richness, with strongest effects emerging for islands. Ants and reptiles, birds and mammals, and vascular plants and spiders form pairs of taxonomic groups with the highest spatial congruence in established alien richness, but drivers explaining richness differ between the taxa in each pair. Across all taxonomic groups, our results highlight the need to prioritize prevention of further alien species introductions to island and coastal mainland regions globally.

Substantial progress has been made in understanding how pathways underlie and mediate biological invasions. However, key features of their role in invasions remain poorly understood, available knowledge is widely scattered, and major frontiers in research and management are insufficiently characterized. We review the state of the art, highlight recent advances, identify pitfalls and constraints, and discuss major challenges in four broad fields of pathway research and management: pathway classification, application of pathway information, management response, and management impact. We present approaches to describe and quantify pathway attributes (e.g., spatiotemporal changes, proxies of introduction effort, environmental and socioeconomic contexts) and how they interact with species traits and regional characteristics. We also provide recommendations for a research agenda with particular focus on emerging (or neglected) research questions and present new analytical tools in the context of pathway research and management.

Prioritization of introduction pathways is seen as an important component of the management of bio-logical invasions. We address whether established alien plants, mammals, freshwater fish and terrestrial invertebrates with known ecological impacts are associated with particular introduction pathways (release, escape, contaminant, stowaway, corridor and unaided). We used the information from the European alien species database DAISIE (www.europe-aliens.org) supplemented by the EASIN catalogue (European Al-ien Species Information Network), and expert knowledge. Plants introduced by the pathways release, corridor and unaided were disproportionately more likely to have ecological impacts than those introduced as contaminants. In contrast, impacts were not associ-ated with particular introduction pathways for invertebrates, mammals or fish. Thus, while for plants management strategies should be targeted towards the appropriate pathways, for animals, management should focus on reducing the total number of taxa introduced, targeting those pathways responsible for high numbers of introductions. However, regardless of taxonomic group, having multiple introduction pathways increases the likelihood of the species having an ecological impact. This may simply reflect that species introduced by multiple pathways have high propagule pressure and so have a high probability of establishment. Clearly, patterns of invasion are determined by many interacting factors and management strategies should reflect this complexity.

© 2021 The Authors., Aims: Understanding fine-grain diversity patterns across large spatial extents is fundamental for macroecological research and biodiversity conservation. Using the GrassPlot database, we provide benchmarks of fine-grain richness values of Palaearctic open habitats for vascular plants, bryophytes, lichens and complete vegetation (i.e., the sum of the former three groups). Location: Palaearctic biogeographic realm. Methods: We used 126,524 plots of eight standard grain sizes from the GrassPlot database: 0.0001, 0.001, 0.01, 0.1, 1, 10, 100 and 1,000 m and calculated the mean richness and standard deviations, as well as maximum, minimum, median, and first and third quartiles for each combination of grain size, taxonomic group, biome, region, vegetation type and phytosociological class. Results: Patterns of plant diversity in vegetation types and biomes differ across grain sizes and taxonomic groups. Overall, secondary (mostly semi-natural) grasslands and natural grasslands are the richest vegetation type. The open-access file ”GrassPlot Diversity Benchmarks” and the web tool “GrassPlot Diversity Explorer” are now available online (https://edgg.org/databases/GrasslandDiversityExplorer) and provide more insights into species richness patterns in the Palaearctic open habitats. Conclusions: The GrassPlot Diversity Benchmarks provide high-quality data on species richness in open habitat types across the Palaearctic. These benchmark data can be used in vegetation ecology, macroecology, biodiversity conservation and data quality checking. While the amount of data in the underlying GrassPlot database and their spatial coverage are smaller than in other extensive vegetation-plot databases, species recordings in GrassPlot are on average more complete, making it a valuable complementary data source in macroecology., GrassPlot development has been supported by the Bavarian Research Alliance (BayIntAn_UBT_2017_58), the Eurasian Dry Grassland Group (EDGG) and the International Association for Vegetation Science (IAVS); IB, CorM, JAC, IGM, DGM, MHe, DL and MTo were supported by the Basque Government (IT936‐16); CorM, IAx, MCh, JDa, PD, MHá, ZL, ZPr, EŠ and LT were supported by the Czech Science Foundation (19‐28491X); TR was supported by the Estonian Research Council (PUT1173); RJP was funded by the Strategic Research Programme of the Scottish Government’s Rural and Environmental Science and Analytical Services Division”; SBa was supported by the GINOP‐2.3.2‐15‐2016‐00019 project; GFi was partially supported by the MIUR initiative “Department of excellence” (Law 232/2016)"; BJA was funded by the Spanish Research Agency (grant AEI/ 10.13039/501100011033); AK, VB, IM, DS, IV and DV were supported by the National Research Foundation of Ukraine (2020.01/0140); MP and AH were supported by the Estonian Research Council (PRG874, PRG609), and the European Regional Development Fund (Centre of Excellence EcolChange); Data collection of HCP was funded by FORMAS (Swedish Research Council for Environment, Agricultural Science and Spatial Planning) and The Swedish Institute; JR was supported by the Czech Science Foundation (grant No. 20‐09895S) and the long‐term developmental project of the Czech Academy of Sciences (RVO 67985939); ATRA was funded by the Grant of Excellence Departments, MIUR‐Italy (ARTICOLO 1, COMMI 314 – 337 LEGGE 232/2016); JMA was supported by Carl Tryggers stiftelse för vetenskaplig forskning and Qatar Petroleum; AAli was supported by the Jiangsu Science and Technology Special Project (Grant No. BX2019084), and Metasequoia Faculty Research Startup Funding at Nanjing Forestry University (Grant No. 163010230), and he is currently supported by Hebei University through Faculty Research Startup Funding Program; ZB was supported by the NKFI K 124796 grant; The GLORIA‐ Aragón project of JLBA was funded by the Dirección General de Cambio Climático del Gobierno de Aragón (Spain); MCs and LDem were supported by DG Environment through the European Forum on Nature Conservation and Pastoralism and Barbara Knowles Fund, in collaboration with Pogány‐havas Association, Romania; JDa was partially supported by long‐term research development project no. RVO 67985939 of the Czech Academy of Sciences; BD and OV were supported by the NKFI KH 126476, NKFI KH 130338, NKFI FK 124404 and NKFI FK 135329 grants; BD, OV and AKe were supported by the Bolyai János Scholarship of the Hungarian Academy of Sciences; BE was funded by the Environmental Department of the Tyrolean Federal State Government, the MAB Programme of the Austrian Academy of Science, the Mountain Agriculture Research Unit and the Alpine Research Centre Obergurgl of Innsbruck University. The GLORIA projects of BE were funded by the EU project no. EVK2‐CT‐2000‐00056, the Earth System Sciences Program of the Austrian Academy of Sciences (project MEDIALPS), the Amt für Naturparke, Autonome Provinz Bozen‐Südtirol, the Südtiroler Wissenschaftsfonds and the Tiroler Wissenschaftsfonds; RGG was supported by the Spanish Ministry of Research to sample GLORIA sites in central Spain (CGL 2008‐00901/BOS) and present works by the Autonomous Region of Madrid (REMEDINAL TE‐CM, S2018/EMT‐4338); MJ was supporteLatviaed by Latvia Grant No. 194051; NP and SŠ were partly supported by the Slovenian Research Agency, core fundings P1‐0403 and J7‐1822.

Aim Species-area relationships (SARs) are fundamental scaling laws in ecology although their shape is still disputed. At larger areas, power laws best represent SARs. Yet, it remains unclear whether SARs follow other shapes at finer spatial grains in continuous vegetation. We asked which function describes SARs best at small grains and explored how sampling methodology or the environment influence SAR shape. Location Palaearctic grasslands and other non-forested habitats. Taxa Vascular plants, bryophytes and lichens. Methods We used the GrassPlot database, containing standardized vegetation-plot data from vascular plants, bryophytes and lichens spanning a wide range of grassland types throughout the Palaearctic and including 2,057 nested-plot series with at least seven grain sizes ranging from 1 cm(2) to 1,024 m(2). Using nonlinear regression, we assessed the appropriateness of different SAR functions (power, power quadratic, power breakpoint, logarithmic, Michaelis-Menten). Based on AICc, we tested whether the ranking of functions differed among taxonomic groups, methodological settings, biomes or vegetation types. Results The power function was the most suitable function across the studied taxonomic groups. The superiority of this function increased from lichens to bryophytes to vascular plants to all three taxonomic groups together. The sampling method was highly influential as rooted presence sampling decreased the performance of the power function. By contrast, biome and vegetation type had practically no influence on the superiority of the power law. Main conclusions We conclude that SARs of sessile organisms at smaller spatial grains are best approximated by a power function. This coincides with several other comprehensive studies of SARs at different grain sizes and for different taxa, thus supporting the general appropriateness of the power function for modelling species diversity over a wide range of grain sizes. The poor performance of the Michaelis-Menten function demonstrates that richness within plant communities generally does not approach any saturation, thus calling into question the concept of minimal area. We thank all vegetation scientists who carefully collected multi‐ scale plant diversity data from Palaearctic Grasslands available in GrassPlot. The Eurasian Dry Grassland Group (EDGG) and the International Association for Vegetation Science (IAVS) sup‐ ported the EDGG Field Workshops, which generated a core part of the GrassPlot data. The Bavarian Research Alliance (grant BayIntAn_UBT_2017_58) and the Bayreuth Center of Ecology and Environmental Research (BayCEER) funded the initial GrassPlot workshop during which the database was established and the cur‐ rent paper was initiated. A.N. acknowledges support by the Center for International Scientific Studies and Collaboration (CISSC), Iran. C.M., I.B., I.G.‐M and J.A.C. were funded by the Basque Government (IT936‐16). D.V. carried out the research supported by a grant of the State Fund For Fundamental Research Ф83/53427. G.F. carried out the research in the frame of the MIUR initiative ‘Department of excellence' (Law 232/2016). I.D. was supported by the Polish National Science Centre (grant DEC‐2013/09/N/NZ8/03234). J.Do. was supported by the Czech Science Foundation (GA 17‐19376S). M.J. was supported by grant by Slovak Academy of Sciences (VEGA 02/0095/19). W.U. ac‐ knowledges support from the Polish National Science Centre (grant 2017/27/B/NZ8/00316).