Irena Axmanová

Veronika Kalusová

Jiří Danihelka

Jürgen Dengler

Jan Pergl

Petr Pyšek

Martin Večeřa

Fabio Attorre

Idoia Biurrun

Steffen Boch

Svetlana Aä

Emiliano Agrillo

Iva Apostolova

Ariel Bergamini

Erwin Bergmeier

Henry Brisse

Laura Casella

Jã¡nos Csiky

Renata Ä †uå¡terevska

Els De Bie

Gianpietro Giusso

Del Galdo

Michele De Sanctis

Iris De Ronde

Panayotis Dimopoulos

Tetiana Dziuba

Úna Fitzpatrick

Xavier Font

Valentin Golub

Friedemann Goral

Ute Jandt

John Janssen

Zygmunt Kä…cki

Mirjana Krstivojeviä

Igor Lavrinenko

Tatiana Lysenko

Corrado Marcenã²

Aaron Pã©rez-Haase

Tomã¡å¡ Peterka

Vadim Prokhorov

Valerijus Raå¡omaviäius

Maria Pilar Rodrã-Guez-Rojo

John S Rodwell

Eszter Ruprecht

Solvita Rå«siå †a

Joachim Schrautzer

Angela Stanisci

Urban Å Ilc

Željko ŠKvorc

Milan Valachoviä

Roberto Venanzoni

Wolfgang Willner

Irena Axmanová¹

Jiří Danihelka¹

Martin Večeřa¹

Timo Conradi

Rosario G Gavilán

Stephan M Hennekens

Jiménez -Alfaro

Ilona Knollová

Anna Kuzemko

Jonathan Lenoir

Artem Leostrin

Jana Medvecká

Erenskjold Moeslund

Dragica Obratov-Petkovic

Jens-Christian Svenning

Ioannis Tsiripidis

Kiril Vassilev

Milan Chytrý¹

Neophyte invasions in European grasslands

Keywords: alien, plant invasion, neophyte, invasion level, invasion success, Europe, European Vegetation Archive (EVA), continental scale, grassland, seminatural vegetation, EUNIS habitat 99 alien, plant invasion, neophyte, invasion level, invasion success, Europe, European Vegetation Archive (EVA), continental scale, grassland, semi-natural vegetation

de niveau recherche, publiés ou non, émanant des établissements d'enseignement et de recherche français ou étrangers, des laboratoires publics ou privés.

Introduction

The human-related spread of alien plants is an important issue worldwide (van Kleunen et al., 2015;[START_REF] Pyšek | Naturalized alien flora of the world: species diversity, taxonomic and phylogenetic patterns, geographic distribution and global hotspots of plant invasion[END_REF] as many alien plant invasions have serious environmental and socioeconomic impacts [START_REF] Vilà | Ecological impacts of invasive alien plants: a meta-analysis of their effects on species[END_REF][START_REF] Pyšek | Scientists' warning on invasive alien species[END_REF]. However, there appears to be significant differences in the levels of alien invasion among regions (Chytrý et al., 2008b;[START_REF] Kalusová | Intercontinental comparison of habitat levels of invasion between temperate North America and Europe[END_REF] and among habitats (Chytrý et al., 2008a;Pyšek et al., 2012a;[START_REF] Kalusová | Alien species pool influences the level of habitat invasion in intercontinental exchange of alien plants[END_REF]. To apply proper risk-assessment schemes and effective management, we need to identify the most vulnerable habitats and the factors responsible for their high levels of alien plant invasion. Europe is an important source of numerous neophytes (i.e. alien plants introduced to a region after 1500 AD) for other continents, but also receives many neophytes from elsewhere [START_REF] Kalusová | Intercontinental comparison of habitat levels of invasion between temperate North America and Europe[END_REF]van Kleunen et al., 2015;2019). The first overviews of alien plants across European countries were based on the data from national and regional floras and additional plant distribution data [START_REF] Weber | The alien flora of Europe: a taxonomic and biogeographical overview[END_REF][START_REF] Lambdon | Alien flora of Europe: species diversity, temporal trends, geographical patterns and research needs[END_REF]. These efforts recently continued with the assessment of naturalized neophytes at the global scale (van Kleunen et al., 2015;2019). However, the data based on regional floras do not necessarily reflect the frequency of neophyte species in different habitats. Therefore, vegetation researchers focused on fine-scale data related to habitat types to get a different point of view of the invasion levels (see [START_REF] Pyšek | Habitat invasion research: where vegetation science and invasion ecology meet[END_REF], for a review). The first comparisons among regions based on vegetation-plot data showed that the same broadly defined habitats had comparable levels of neophyte invasions (Chytrý et al., 2008b). Despite different absolute invasion levels, the most and least invaded habitats were the same when comparing European and North American regions [START_REF] Kalusová | Intercontinental comparison of habitat levels of invasion between temperate North America and Europe[END_REF], suggesting that habitat typologies are a key factor to understand plant invasions. Generally, the most invaded habitats seem to be human-made habitats (e.g. arable fields, wastelands), while natural vegetation such as high-altitude grasslands or mires are less invaded (Chytrý et al., 2008b;[START_REF] Lambdon | Alien flora of Europe: species diversity, temporal trends, geographical patterns and research needs[END_REF]. The recently compiled European Vegetation Archive (EVA; [START_REF] Chytrý | European Vegetation Archive (EVA): An integrated database of European vegetation plots[END_REF] enabled even deeper insights into the neophyte invasion patterns at the level of fine-scale habitats, already reported for European woodlands [START_REF] Wagner | Alien plant invasions in European woodlands[END_REF] and coastal dunes [START_REF] Giulio | Alien flora across European coastal dunes[END_REF]. However, neophyte invasions across grassland habitats have not been studied so far at the European extent. After forests, grasslands represent the second most widespread broad vegetation type in Europe, occurring across large gradients of soil moisture and pH, nutrient availability and climate. Grassland habitats also bear the imprint of the past, not only glacial-interglacial cycles and postglacial migration, but also traditional management practices [START_REF] Pärtel | Biodiversity in temperate European grasslands: Origin and conservation[END_REF][START_REF] Feurdean | Biodiversity-rich European grasslands: Ancient, forgotten ecosystems[END_REF]. All these factors shape current grassland vegetation and result in high variability of vegetation types [START_REF] Chytrý | EUNIS Habitat Classification: expert system, characteristic species combinations and distribution maps of European habitats[END_REF]. Previous comparisons of broadly defined habitat types showed that different levels of alien plant invasion could be explained by several factors, especially resource availability, propagule pressure and climate (Chytrý et al., 2008a). Another important aspect affecting alien plant invasions are disturbances [START_REF] Hobbs | Disturbance, diversity and invasion: Implications for conservation[END_REF] 5 human-made habitats and alluvial forests are generally occurring in the most fertile parts of the countries and are subjected to repeated introductions of neophytes, grasslands are likely to be less exposed to alien plant propagule pressure [START_REF] Botham | Do urban areas act as foci for the spread of alien plant species? An assessment of temporal trends in the UK[END_REF]. The type, frequency and intensity of disturbance in grasslands differ from those typical of human-made habitats or forests. Consequently, also the geographic patterns of neophyte invasions in grasslands and forests differ considerably (Divíšek and Chytrý, 2018). Therefore, in addition to the recent assessment of neophyte invasions across European woodlands [START_REF] Wagner | Alien plant invasions in European woodlands[END_REF], a reliable assessment of neophyte invasions across European grasslands is also needed. To meet this aim, we used an up-to-date version of the largest European vegetation-plot database (EVA; [START_REF] Chytrý | European Vegetation Archive (EVA): An integrated database of European vegetation plots[END_REF] to provide a synthetic overview of neophyte invasions in European grassland vegetation, asking the following questions: (i) How do invasion levels change across European countries? (ii) Which grassland types are the most invaded? (iii) Which alien species are the most successful invaders of European grasslands?

Methods

Region definitions

Regions used in our study mostly correspond to countries, but some large islands or archipelagoes with distinct biogeographic position were treated as separate regions (Baleares, Corsica, Crete, Sardinia, and Sicily with Malta). We also followed the region delimitation concept of the Euro+Med PlantBase (2006)(2007)(2008)(2009)(2010)(2011)(2012)(2013)(2014)(2015)(2016)(2017)(2018)(2019)(2020), and divided European Russia into seven regions (Russia North, Russia Northwest, Russia Kaliningrad, Russia Central, Russia South, Russia Caucasus and Russia East). For the list of all 49 studied regions and the detailed number of plots per region, see Appendix S3.

Dataset preparation and filtering

We used 465,629 grassland vegetation plots provided by the European Vegetation Archive (EVA; [START_REF] Chytrý | European Vegetation Archive (EVA): An integrated database of European vegetation plots[END_REF]. The plots were classified to European Nature Information System grassland habitat types at the third level of the classification hierarchy (hereafter EUNIS habitats or habitats) using the EUNIS-ESy expert system [START_REF] Chytrý | EUNIS Habitat Classification: expert system, characteristic species combinations and distribution maps of European habitats[END_REF] in the JUICE software (Tichý, 2002). Apart from grasslands in a narrow sense (EUNIS group R), we also included coastal grasslands of grey dunes, which belong to EUNIS group N (coastal habitats). An overview of contributing databases and numbers of plots assigned to particular EUNIS habitat types is available in Appendices S1 and S2 in the Supplementary Materials. We also merged these habitats to broader habitat types (habitat groups, see Appendix S2 for details on merging), enabling us to detect more general patterns in analyses and comparisons at a coarser scale. Data from the National Vegetation Database of Denmark could not be classified by the EUNIS-ESy expert system, because they lack records of cover-abundances of species. Therefore, we calculated the Bray-Curtis dissimilarity index to find the most similar plots from other countries with known classification and assigned the Danish plots to the same habitat groups 6 (all plots with similarity lower than 15%, i.e., Bray-Curtis dissimilarity > 0.85, were excluded; in total 1,885 vegetation plots were assigned to habitat groups). For our analyses, we filtered vegetation plots using the following criteria. First, we removed: (i) plots that could not be assigned to any EUNIS grassland habitat type at the third level of classification (transitional between different grassland habitat types); (ii) plots with a cover of woody species (trees or shrubs taller than 0.5 m) higher than 10%; (iii) plots dominated by Pteridium aquilinum (classified as R54 in EUNIS habitat); and (iv) plots from forest clearings (R57). Second, we excluded plots without geographic coordinates or with a location uncertainty ≥ 7000 m. Third, we selected only plots with a surface area ranging between 10 and 100 m 2 to limit the area effect (85% of the plots in original selection were of this size). However, for Nordic countries we retained plots between 1 and 100 m 2 , since the small-size plots traditionally prevail there. We also kept plots with unknown size (based on the plot size histogram, we assumed most of the unknown sizes were within the range of 10-100 m 2 ). Fourth, we kept only the vegetation plots sampled between 1970 and 2018 to lower the effect of vegetation changes across time. Fifth, to limit potential pseudo-replication due to spatial autocorrelations, we used geographical position and compositional similarity to define group of plots that need resampling (we selected all plots with similarity of 80% or more, i.e. ≤ 0.2 of Bray-Curtiss dissimilarity index) sampled within a distance of less than 1 km). From each pair of plots with highest similarity, we randomly selected just one and if needed repeated the resampling. Sixth, to further reduce over-representation of some regions in the dataset, we calculated the density of plots belonging to each habitat type for each region. Subsequently, the corresponding density threshold defining outlier values (upper quartile + 1.5 × interquartile range) was calculated for each habitat type. This threshold was used as a limit for the maximum number of plots to be retained for a particular habitat in a given region. The plots to fit this limit were randomly selected. After filtering, the final dataset included 97,417 plots.

Species status assignment

We considered only vascular plants in our study, including trees and shrubs. The nomenclature was unified to follow Euro+Med (2006-2020) PlantBase concept, or The Plant List (TPL, 2013) if the taxon was not included in Euro+Med. We assigned the native, alien or uncertain status to all the plant taxa with respect to the region where they were recorded. We included archaeophytes among native species, hence the alien plants in our study corresponded to neophytes only (i.e., alien species that arrived in the region after 1500 AD, see Pyšek and Jarošík, 2005). Archaeophytes were not treated separately as their lists are lacking or incomplete in many countries, and in many cases their status as arquaeophyte or native in a given country or region is doubtful. If a species was considered native in at least one part of a region, we assigned its native status for that region. We assigned the native/alien status to species using the Euro+Med PlantBase (2006)(2007)(2008)(2009)(2010)(2011)(2012)(2013)(2014)(2015)(2016)(2017)(2018)(2019)(2020), [START_REF] Daisie | Handbook of alien species in Europe[END_REF], the GLONAF database (van Kleunen et al., 2019), national and regional species checklists and knowledge of local experts. All sources are listed in Appendix S4. Taxa for which we could not decide their status were excluded (mostly genus-level records). For a status assignment at the plot level, we

F o r R e v i e w O n l y
7 used intraspecific data, as sometimes one subspecies can be native while another subspecies is considered as neophyte. For comparison between species lists of regions and habitats, we summarised the results at the species level (considering the subspecies status assignment) to make our results comparable with recent studies of other European habitats (e.g., [START_REF] Wagner | Alien plant invasions in European woodlands[END_REF][START_REF] Giulio | Alien flora across European coastal dunes[END_REF]. We further classified neophytes into groups according to their origin: A, neophyte species originating from outside of Europe (Africa, America, Asia, Australia); E, species native to some parts of Europe but considered to be neophytes elsewhere in Europe, C, other origins, i.e., anecophytes or species with hybrid origin (see also Wagner et al., 2017 for details). We also classified all neophytes with respect to their prevailing life form: phanerophytes, chamaephytes, hemicryptophytes, geophytes, therophytes, hydrophytes and woody lianas. For the list of sources used for this assignment, see Appendix S5.

Data analyses

We assessed the levels of invasions using different metrics (see also [START_REF] Catford | Quantifying levels of biological invasion: towards the objective classification of invaded and invasible ecosystems[END_REF][START_REF] Wagner | Alien plant invasions in European woodlands[END_REF][START_REF] Giulio | Alien flora across European coastal dunes[END_REF]: (1) percentage of neophytes in the grassland species list, (2) frequency of neophytes across all occurrences (sum of individual plot records) of all species in a given subset, (3) relative frequency of invaded plots (i.e., plots with at least one neophyte present), (4) mean and median of relative species richness of neophytes per plot in a given subset (relative species richness of neophytes refers to the number of neophytes/all species in the plot *100), (5) relative neophyte cover per plot in a given subset (relative to total cover in the plot). To compare the species lists of habitat groups with different numbers of plots, we selected only neophytes represented in at least 0.5% and in at least 1% of the plots of that habitat group. Note that we selected such low levels of threshold because most of the plots were free of neophytes. The differences in relative neophyte richness and relative neophyte cover among habitat groups were tested using the Kruskal-Wallis multiple comparison test in the package pgirmess [START_REF] Giraudoux | Miscellaneous functions for analysis and display of ecological and spatial data. Package "pgirmess[END_REF] in the R software (R Core Team, 2012). Total plant cover and neophyte cover were calculated as a sum of individual plant covers considering their possible overlap [START_REF] Fischer | On the combination of species cover values from different vegetation layers[END_REF]. To track geographical patterns in the level of invasions, we assigned vegetation plots into the UTM grid cells spanning 50 km × 50 km. For each grid, we calculated the relative frequency of invaded plots, i.e., what percentage of plots within a given grid cell have at least one neophyte present. We prepared separate maps with respect to habitat groups and different origin categories of neophytes using the R packages 'raster' [START_REF] Hijmans | raster: Geographic Data Analysis and Modeling[END_REF], 'rgdal' [START_REF] Bivand | rgdal: Bindings for the 'Geospatial' Data Abstraction Library[END_REF], 'spatialEco' [START_REF] Evans | spatialEco: Spatial Analysis and Modelling Utilities[END_REF] and 'berryFunctions' [START_REF] Boessenkool | berryFunctions: Function Collection Related to Plotting and Hydrology[END_REF]. We mapped only grid cells containing more than ten plots, in order to avoid interpreting random patterns. The data were processed using the R software v.3.6.0 (R Core Team, 2019).

Results

Our final grassland dataset included in total 8,212 species, out of which only 536 were classified as neophytes (approximately 6.5% of all species) with almost the same number of neophytes originating from Europe (3.0%) and outside of Europe (3.1%). Most neophytes in grasslands were present in very few plots (median of 2 plots), while only three species, namely Onobrychis viciifolia, Erigeron annuus and E. canadensis, were recorded in more than 1% of all plots (Table 1). When comparing the relative frequency of species occurrences in plots, European neophytes correspond to 0.17% (4,259 of all 2,565,674 occurrences), while those from outside of Europe to 0.33% (8,473), which is different from percentages based on the grassland species list (see above, or Figure 1). The prevailing life forms of neophytes in our study were hemicryptophytes (32%) and therophytes (31%), followed by phanerophytes (21%) (Figure 2). Most of the vegetation-plot records included only native species and archaeophytes, while invaded plots, i.e., plots with at least one neophyte, represented only 11.9% of all plots (7.1% of plots if only non-European neophytes were considered, Appendix S6). The proportion of invaded plots varied greatly among individual regions and habitat types. The means of relative neophyte species richness per plot also differed, although the medians were close or equal to zero, suggesting the prevalence of neophyte-free plots. Summary statistics can be found in Appendix S6 and S7 for regions and in Appendix S8 and S9 for habitat groups and EUNIS habitats. A list of neophytes recorded per regions is available in Appendix S10. We did not reveal any distinct geographical pattern in the distribution of neophytes across Europe (see Figure 3 for overall pattern and other maps in Appendix S12 for separate habitat groups or neophytes according to their origin). Higher invasion levels can be seen in northern European countries, where this is mostly due to the higher number of European neophytes (see also Appendix S6). In addition, there is a slightly higher tendency for invaded plots to occur in coastal areas. When comparing only the relative frequency of non-European neophytes, the regions with the highest representation of neophytes relative to all species were Poland (3.9%), Netherlands (3.8%), Lithuania (3.5%), Denmark (3.0%) and Germany (3.0%), for more see Appendix S6. Analyses at the third hierarchical level of EUNIS habitat classification show that sandy habitats (both inland and coastal) have the highest frequency of invaded plots across different types and geographical regions (on average around 20%, with a maximum of 31%). Comparable percentages of invaded plots were also found in lowland wet tall-herb grasslands, in both temperate and Mediterranean types. In contrast, high-altitude grasslands, especially alpine and oromediterranean, had the lowest percentages of invaded plots (Figure 4). Comparison among the habitat groups revealed that most neophyte species (90% when considering only species recorded in at least five plots) occur in more than one habitat group , indicating their low degree of specialization (see Table 2 for the most frequent neophytes per habitat group). The highest numbers of neophyte species were recorded

F o r R e v i e w O n l y
9 in mesic, wet and dry grasslands, followed by sandy grasslands (Table 3). However, removing rare species with few occurrences (recorded in less than 0.5 or 1% of the plots from the respective habitat) from these counts lowers the number of neophyte species considerably (Figure 5a,c). After the exclusion of rare neophytes, sandy, wet and mesic habitats harboured the highest cumulative numbers of neophyte species, while the oromediterranean and alpine grasslands the lowest. A similar pattern can also be seen in the relative frequency of invaded plots per habitat group (Figure 5b,d). Note that in dry grasslands, the relative frequency of invaded plots reaches almost the same percentages as in mesic and wet grasslands, but the number of neophytes in dry grassland species list is much lower, suggesting their higher frequency in the plots. While rocky, dry and mesic habitat groups have a higher number of European neophytes than those of non-European origin, sandy, wet and saline habitats showed a reverse pattern. We revealed the highest prevalence of non-European neophytes in wet grasslands (Table 3). For mean relative neophyte richness and other summary statistics calculated across plots, see Table 4. Kruskal-Wallis multiple comparisons of both the mean relative neophyte richness and the mean relative neophyte cover per habitat groups confirmed the above described pattern (

Discussion

Geographical patterns

We found a relatively low proportion of neophyte species in European grasslands (only 6.5% of all species in the dataset). Such low levels of invasion are similar to values reported from European forests [START_REF] Wagner | Alien plant invasions in European woodlands[END_REF] and coastal vegetation [START_REF] Giulio | Alien flora across European coastal dunes[END_REF], but they are in contrast to those reported from North-American habitats [START_REF] Lonsdale | Global Patterns of Plant Invasions and the Concept of Invasibility[END_REF][START_REF] Kalusová | Intercontinental comparison of habitat levels of invasion between temperate North America and Europe[END_REF]. The several times higher numbers of neophytes in North America, even when comparing similar grasslands (15% reported from prairies by [START_REF] Stohlgren | Exotic Plant Species Invade Hot Spots of Native Plant Diversity[END_REF], support the view that Europe is donating more alien species than it is receiving in exchange [START_REF] Seastedt | Mechanisms of plant invasions of North American and European grasslands[END_REF][START_REF] Stohlgren | Widespread plant species: natives versus aliens in our changing world[END_REF][START_REF] Hejda | Native-range habitats of invasive plants: are they similar to invaded-range habitats and do they differ according to the geographical direction of invasion?[END_REF], which holds also specifically for grasses [START_REF] Monnet | Historical legacies and ecological determinants of grass naturalizations worldwide[END_REF]. We did not find any distinct geographical pattern, although there were higher percentages of invaded plots in northern Europe (mostly due to a higher abundance of neophytes of European origin there) and in coastal habitats. The neophytes of European origin were mostly from the Mediterranean, i.e. native to southern Europe from where they spread to suitable habitats in other regions. Depending on when they reached individual countries, the same species can be classified as either an archaeophyte or neophyte (Pyšek et al., 2005). Moreover, plant ranges do not reflect only recent climate but also the landscape history, especially Pleistocene glaciations, as the possibilities of postglacial migration towards the north were limited [START_REF] Normand | Postglacial migration supplements climate in determining plant species ranges in Europe[END_REF]. A large number of otherwise frequent European species have probably been able to colonize northern countries only as a consequence of human-induced changes of habitats [START_REF] Lambdon | Alien flora of Europe: species diversity, temporal trends, geographical patterns and research needs[END_REF]. The shares of non-European and European neophytes were almost equal in terms of numbers of species occurring in grassland, but the latter group prevailed when considering the relative frequency of occurrences across the plots (accounting for 58.6% of all neophyte occurrences, most of them from North America). This is in agreement with greater representation of species from other continents reported by previous large-scale European studies [START_REF] Wagner | Alien plant invasions in European woodlands[END_REF][START_REF] Giulio | Alien flora across European coastal dunes[END_REF]. Such pattern results mostly from the fact that there are fewer regions in Europe where species of European origin can be recorded as neophytes, although alternative explanations can also be raised (see [START_REF] Wagner | Alien plant invasions in European woodlands[END_REF]. We found the highest levels of invasion (relative number of neophytes in the species list) in Poland, the Netherlands, Lithuania, Denmark and Germany, where both European and non-European neophytes were present in relatively high numbers. We expect that in these countries, there was both the effect of glaciation and at the same time the presence of coastal habitats which are generally more invaded (Chytrý et al., 2008b;Dawson et al. 2017;[START_REF] Giulio | Alien flora across European coastal dunes[END_REF]. Another important factor supporting higher invasion levels can also be the recent changes in traditional management (e.g. Timmermann et al., 2015). We note that natural factors such as grazing by wild herbivores also maintained a presence of grasslands in temperate Europe before agriculture and pastoralism or even before the arrival of Homo sapiens to the continent [START_REF] Svenning | A review of natural vegetation openness in north-western Europe[END_REF][START_REF] Feurdean | Biodiversity-rich European grasslands: Ancient, forgotten ecosystems[END_REF]. However, nowadays many grasslands experience both

F o r R e v i e w O n l y
11 reduced wild grazing and reduced management, leading to secondary succession and vegetation changes that can support the success of neophytes (e.g. Swacha et al., 2018).

Habitat comparison

We confirmed that grasslands generally have low levels of invasion, which is in contrast to highly invaded human-made habitats, alluvial forests and herb-dominated river fringes (Chytrý et al., 2008b;[START_REF] Kalusová | Intercontinental comparison of habitat levels of invasion between temperate North America and Europe[END_REF]. However, although the overall frequency of invaded plots was rather low (less than 12% of all plots), EUNIS habitat types varied from 0 to 31% invaded plots. The highest percentages of invaded plots were found in sandy habitats and in lowland tall-herb wet meadows, while grasslands at high altitudes were almost free of any neophytes. Such pattern is in accordance with conclusions of earlier studies that neophytes preferably invade habitats with sufficient resources, under milder climate and that they benefit from disturbances that suppress native species (Chytrý et al., 2008a;[START_REF] Pyšek | Habitat invasion research: where vegetation science and invasion ecology meet[END_REF]. When comparing neophytes among habitat groups, we did not find any typical habitat specialists. Similarly, also [START_REF] Giulio | Alien flora across European coastal dunes[END_REF] concluded that most (94%) of the neophytes found in their study of coastal dunes were generalists occurring also in other habitats. Chytrý et al. (2008b) revealed comparable habitat levels of invasions among regions, although the species lists of neophytes were different, suggesting that the habitat properties are more important than geographical position within Europe. Since sandy and wet grasslands differ in many habitat characteristics, we can also expect different strategies of neophytes invading them. Life-form comparison of neophytes revealed relatively high representation of two distinct groups in our grassland dataset species list, namely therophytes and hemicryptophytes (each group represented around 30%), which supports our expectation of different strategies. Wet grasslands mostly have high total cover, and gaps occur only rarely after disturbances; thus, competitive hemicryptophytes are more successful. In contrast, the unstable soil in sandy habitats does not allow the formation of closed stands, and many open gaps are available to support therophytes. The spread of these short-lived, ruderal species can be recently detected also in other vegetation types, such as dry grasslands. As present weather fluctuations and drought events are leading to decreased cover or even disappearance of some native species, new gaps emerge in the stands and therefore also opportunities for new species to successfully germinate [START_REF] Fischer | Weather fluctuations drive short-term dynamics and long-term stability in plant communities: A 25-year study in a Central European dry grassland[END_REF]. Although canopy openness seems to be important, it does not ensure the success of the neophytes alone. We can expect that the rather open stands of alpine and oromediterranean habitats are so extreme and nutrient-poor that only species adapted to such conditions can grow there, which limits the number of suitable neophytes (Chytrý et al., 2008b). Compared to lowlands, grasslands in higher altitudes historically experienced lower human impact and intensity of disturbances, thus suppressing or at least reducing success of neophytes (see also [START_REF] Medvecká | Fifty years of plant invasion dynamics in Slovakia along a 2,500 m altitudinal gradient[END_REF];

F o r R e v i e w O n l y 12

Successful neophytes

Neophytes themselves significantly differ in their ability to overcome abiotic and reproductive barriers and establish sustainable populations in new environments. Some species are able to invade only the same or similar habitat from which they originated and to which they are adapted. Therefore, the more habitats the species occupies in its native range, the higher are the chances that it will become a successful invader [START_REF] Hejda | Native-range habitats of invasive plants: are they similar to invaded-range habitats and do they differ according to the geographical direction of invasion?[END_REF][START_REF] Kalusová | Naturalization of European plants on other continents: The role of donor habitats[END_REF], and the most successful species that reach the invasion stage extend their habitat niche in the invaded regions, compared to the region of origin [START_REF] Hejda | Invasion success of alien plants: do habitats affinities in the native distribution range matter?[END_REF]. The association with human-made habitats and disturbances in the native range also seems to support invasion success [START_REF] Kalusová | Naturalization of European plants on other continents: The role of donor habitats[END_REF]. This is in agreement with our findings that many neophytes in our grassland dataset were not habitat specialists but short-lived ruderal species, using opportunities of establishing in open gaps after disturbances. Divíšek et al. (2018) suggested that while having similar traits as other species within the community enables naturalization in that particular habitat, a difference in at least some traits (such as plant height) allows the neophyte species to prevail in the community and become invasive. This points at another neophyte group -species growing in relatively productive sites, with intense competition for resources (e.g. Solidago spp., Symphyotrichum spp.). Within our dataset, we found mostly neophytes with few occurrences only. Despite being rare in semi-natural grasslands, it is probable that many of these species are more abundant in more suitable, human-made habitats of the same regions. We also have to note that many neophytes are relatively new in Europe, and longer residence time is probably needed for them to become more widespread (Pyšek and Jarošík 2005). The most frequent neophyte in our dataset was Onobrychis viciifolia, a perennial species native to southern and southeastern Europe and western Asia, from where it was intentionally introduced to the rest of Europe (but also to North America and Australia) as a forage crop [START_REF] Mora-Ortiz | Onobrychis viciifolia; a comprehensive literature review of its history, etymology, taxonomy, genetics, agronomy and botany[END_REF]. After the first introductions, it started to spread spontaneously to suitable habitats. The second most frequent species was Erigeron canadensis, an annual or overwintering species native to southern Canada and having a broad climatic tolerance. Although it is mostly found in human-made habitats (orchards, vineyards, roadsides, arable fields, ruderal sites), it has broad climatic tolerance and is adaptable to different environments. Thanks to very effective wind dispersal, it can quickly establish in open gaps after disturbance and start to grow rapidly [START_REF] Weaver | The biology of Canadian weeds. 115. Conyza canadensis[END_REF]. It was reported as the most successful alien species also in European coastal dunes [START_REF] Giulio | Alien flora across European coastal dunes[END_REF] and across Europe in general [START_REF] Lambdon | Alien flora of Europe: species diversity, temporal trends, geographical patterns and research needs[END_REF]. The third most frequent species was Erigeron annuus, an annual species with prevailing apomictic reproduction, native to North-American tall-grass prairies. This species also has effective wind dispersal and is widespread across Europe. Although it has been shown that its frequency declines towards higher altitudes, growth performance and seed production are comparable with lowland populations, and the reason for the decline is probably its greater winter mortality (Trtikova et al. 2011). In contrast to Onobrychis, the spread of which was directly supported by humans, both Erigeron species

F o r R e v i e w O n l y
13 seem to be so successful mostly due to high seed production and long-distance seed dispersal [START_REF] Weaver | The biology of Canadian weeds. 115. Conyza canadensis[END_REF] as high propagule pressure is one of the key factors in neophyte invasions [START_REF] Pyšek | Habitat invasion research: where vegetation science and invasion ecology meet[END_REF].

Dataset limitations

There are several possible limitations of our data. Firstly, EVA database [START_REF] Chytrý | European Vegetation Archive (EVA): An integrated database of European vegetation plots[END_REF] comprises data originally sampled for phytosociological surveys; therefore, we can expect that they largely originate from preferential sampling [START_REF] Michalcová | Bias in vegetation databases? A comparison of stratifiedrandom and preferential sampling[END_REF], as scientists studying natural and semi-natural vegetation often show tendency to avoid plots with alien plants. Therefore, only when the neophyte is more abundant in the landscape, it is also recorded in the plots. Secondly, when analyzing these data, we cannot easily track the changes in vegetation following the abandonment of traditional management, although this seems to be an important factor. For example, if a grassland site is severely disturbed and the management is no longer applied, the succession might lead towards ruderal sites dominated by invasive neophytes (such as Solidago gigantea), which are no longer classified as grasslands and included in our dataset.

Finally, yet importantly, the data coverage across regions is unequal, and although we stratified the dataset to avoid oversampling of some regions, there were regions less well represented in the data. We would also like to point at the questionable status of some species, which can affect the overall patterns (but see [START_REF] Courchamp | Invasion biology and uncertainty in native range definitions: response to Pereyra 2019[END_REF]. As most problematic we see grasses, which were traditionally supported by humans and are hard to distinguish in archaeological findings, such as Lolium multiflorum, Festuca brevipila, but most notably Arrhenatherum elatius. Although some sources suggest neophyte status of Arrhenatherum (see for example [START_REF] Poschlod | Geschichte der Kulturlandschaft: Entstehungsursachen und Steuerungsfaktoren der Entwicklung der Kulturlandschaft[END_REF], based on previous archaeological findings we accepted it as an archaeophyte (see Pyšek et al., 2012b). In the opposite case, it would be one of the most successful neophyte across Europe.

Conclusions

We provided the first overview of neophyte invasion patterns in grassland vegetation across Europe based on the most comprehensive dataset of vegetation plots existing to date. Although we found generally low invasion levels, there were distinct differences between individual habitat types. While sandy habitats and lowland wet grasslands were the most invaded, high-altitude grasslands were almost free of neophytes. A high proportion of neophytes invading grasslands are therophytes with broad habitat niches.

F o r R e v i e w O n l y 14
We thank Lubomír Tichý for help with the assignment of the Danish vegetation plots into broad vegetation types; Petra Štěpánková for help with life-form assignment; Viktoria Wagner for help with R scripts; Jan Divíšek for help with geographical stratification; Svetlana Aćić, Emiliano Agrillo, Iva Apostolova, Ariel Bergamini, Erwin Bergmeier, Henry Brisse, Laura Casella, János Csiky, Renata Ćušterevska, Els De Bie, Gianpietro Giusso del Galdo, Michele De Sanctis, Iris de Ronde, Panayotis Dimopoulos, Tetiana Dziuba, Úna FitzPatrick, Xavier Font, Valentin Golub, Friedemann Goral, Ute Jandt, John Janssen, Zygmunt Kącki, Mirjana Krstivojević Ćuk, Igor Lavrinenko, Tatiana Lysenko, Corrado Marcenò, Aaron Pérez-Haase, Tomáš Peterka, Vadim Prokhorov, Valerijus Rašomavičius, Maria Pilar Rodríguez-Rojo, John S. Rodwell, Eszter Ruprecht, Solvita Rūsiņa, Joachim Schrautzer, Angela Stanisci, Urban Šilc, Željko Škvorc, Milan Valachovič, Roberto Venanzoni, Wolfgang Willner and Sergey Yamalov for data contributions from vegetation-plot databases they manage. Last but not least, we thank all the people who collected the plot data on grassland vegetation in the field. 1. A list of the most frequent neophyte species recorded in European grasslands assessed by (a) number of plots across all regions where the species is considered neophyte (from a total of 97,417 plots) and (b) number of regions where the species occurred as a neophyte (from a total of 49; for more details on regions, see Appendix S6, S7 and S10). E -species with origin within Europe (they are considered neophytes only in some European regions, and their occurrences in the regions with native status were not counted), A -non-European origin, C -other origin (anecophytes, hybrids). 2. A list of the most frequent neophyte species per habitat group assessed by (a) number of plots across all regions where the species is considered to be a neophyte and (b) number of regions where the species is considered to be a neophyte. The group of oromediterranean habitats (recorded in six regions) is excluded here due to the absence of neophytes. All the neophyte species in the group of alpine habitats were recorded in one region only; therefore, they cannot be sorted by the number of regions in (b). Origin 'A' refers to non-European origin, 'E' to European origin and 'C' to another origin (anecophytes, hybrids). For details on included intraspecific taxa see Table 1.

F o r R e v i e w O n l y F o r R e v i e w O n l y F o r R e v i e w O n l y F o r R e v i e w O n l y F o r R e v i
(a) (b)

Habitat group (plots) 4. Summary statistics for habitat groups: mean, standard error of the mean (SEM) and median of species richness and relative abundance of neophyte species calculated across plots. Oromedit. refers to oromediterranean habitat group. Species richness refers to all species recorded in the plot. Neophytes include all neophyte species irrespective of their origin, while they are separated in the latter categories. Kruskal-Wallis test shows significant differences at the level of p= 0.01.

0 0 0 0 0 0.1 median 0 0 0 0 0 0 0 0 Non-European origin [%] mean 1.8 0.1 0.1 0 0.4 0.8 0 0.7 SEM 0.1 0 0 0 0 0 0 0.1 median 0 0 0 0 0 0 0 0 European origin [%] mean 0.4 0.2 0.2 0 0.2 0.1 0 0.1 SEM 0 0 0 0 0 0 0 0 median 0 0 0 0 0 0 0 0 Other [%] mean 0 0 0 0 0.1 0.1 0 0.1 SEM 0 0 0 0 0 0 0 0 median 0 0 0 0 0 0 0 0 Kruskal-

Figure 1 .

 1 Figure1. Absolute and relative numbers of native (including archaeophytes) and neophyte species in European grasslands. Representation in the grassland species list (a) considers native only those species that are in all countries included in the native category, while species of European origin considered as a neophyte in at least one country were included in the neophyte numbers. In total, 242 species had double status within Europe (approximately 2.9 %). Overview based on all occurrences (b) considers each record of plant species in a plot separately (see Appendix S10 for details).

Figure 2 .

 2 Figure 2. Life forms of neophytes in European grasslands.

Figure 3 .

 3 Figure 3. Relative frequency of invaded plots calculated per UTM 50 × 50 km grid cells. Colour-scale categories were defined to reflect the distribution of values (breaks calculated by k-means). Grey colour indicates cells with only notinvaded plots (0 %). Cells with less than ten plots are not mapped. Maps for habitat groups and neophyte origin categories are in Appendix S12.

Figure 4 .

 4 Figure 4. Relative frequency (%) of plots with neophytes in individual EUNIS habitats. Corresponding habitat groups are indicated by the labels to the right part of the bars. Note that last nine habitats (starting with R17) do not have any invaded plots and are sorted alphabetically by their EUNIS codes. For other statistics, see Appendix S8 and S9. Unclassified plots from Denmark are not included.

Figure 5 .

 5 Figure 5. Overview of the species lists of neophytes by habitat groups. (a) Neophytes occurring in at least 0.5 or 1% of plots, respectively; (b) Relative frequency of plots invaded by neophytes; (c) Non-European neophytes occurring in at least 0.5 or 1% of plots; (d) Relative frequency of plots invaded by non-European neophytes.

Table 4 ,

 4 Appendix S11).

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  List of neophytes recorded per region. Appendix S11 Comparison of relative neophyte species richness and neophyte cover among habitat groups. Appendix S12 Maps of the proportion of invaded plots in individual habitats relative to the neophyte origin categories.

	Appendix S10 F o
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	19 20

Table

  Xanthium orientale (subsp. orientale, subsp. californicum, subsp. italicum, subsp. riparium). Oenothera biennis agg. refers to a group of taxa which are difficult to determine and usually not distinguished.

								No. of
			No. of					regions
	Taxon	Origin	plots	%		Taxon	Origin	(max. 49)
	Onobrychis viciifolia	E	1442	1.5		Erigeron canadensis	A	27
	Erigeron annuus*	A	1312	1.3		Medicago sativa*	C	22
	Erigeron canadensis	A	1188	1.2		Oenothera biennis agg.	A	20
	Trifolium hybridum*	C	794	0.8		Erigeron annuus*	A	19
	Medicago sativa*	C	743	0.8		Matricaria discoidea	A	19
	Solidago gigantea Epilobium ciliatum* Matricaria discoidea Juncus tenuis	A A A F o A r 609 408 348 323	0.6 0.4 0.4 0.3		Xanthium orientale* Juncus tenuis Bidens frondosus Onobrychis viciifolia	A A A E	19 18 17 17
	Impatiens glandulifera Oenothera biennis agg. Solidago canadensis Senecio leucanthemifolius* E A A A Impatiens parviflora A Lolium multiflorum E	304 244 R 241 e 212 210 v 0.3 0.3 0.2 0.2 0.2 i 208 0.2 e	Veronica persica Robinia pseudoacacia Epilobium ciliatum* Impatiens glandulifera Solidago canadensis Impatiens parviflora	A A A A A A	17 16 15 15 15 14
	Bidens frondosus Berteroa incana	A E	199 186	0.2 0.2 w Solidago gigantea Amaranthus retroflexus	A A	14 13
	Veronica persica Achillea ptarmica Xanthium orientale*	A E A	178 171 167	0.2 0.2 0.2	Galinsoga quadriradiata Lolium multiflorum Oxalis stricta O n	A E A	12 12 12
						l
						y
				21

*Taxon names marked by asterisk include the following subspecies: Epilobium ciliatum (subsp. ciliatum, subsp. adenocaulon), Erigeron annuus (subsp. annuus, subsp. septentrionalis), Medicago sativa (subsp. sativa, nothosubsp. varia), Senecio leucanthemifolius subsp. vernalis, Trifolium hybridum (subsp. hybridum, subsp. elegans),

Table 3 .

 3 Overview of the species-pool data and relative frequency of invaded plots (plots with the presence of at least one neophyte) for habitat groups. Oromedit. refers to oromediterranean habitat group.

		Taxon	Origin	No. of plots	%	Taxon	Origin	No. of regions	(max)
	Sandy	Erigeron canadensis	A 492	11.3	Erigeron canadensis	A	20 (38)
	(4359)	Oenothera biennis agg.	A 148	3.4	Oenothera biennis agg.	A	10 (38)
		Senecio leucanthemifolius	E 113	2.6	Xanthium orientale	A	8 (38)
		Xanthium orientale	A 79	1.8	Robinia pseudoacacia	A	7 (38)
		Berteroa incana	E 71	1.6	Ambrosia artemisiifolia	A	6 (38)
	Rocky	Berteroa incana	E 42	0.9	Erigeron canadensis	A	10 (33)
	(4671)	Erigeron canadensis	A 36	0.8	Onobrychis viciifolia	E	7 (33)
		Onobrychis viciifolia	E 36	0.8	Erigeron annuus	A	5 (33)
		Dianthus caryophyllus	E 34	0.7	Medicago sativa	C	5 (33)
		Oxalis dillenii	A 20	0.4	Oenothera biennis agg.	A	4 (33)
	Dry	Onobrychis viciifolia	E 1073	4.2	Erigeron canadensis	A	17 (43)
	(25430) Erigeron annuus	A 307	1.2	Medicago sativa	C	17 (43)
		Medicago sativa	C 272	1.1	Erigeron annuus	A	16 (43)
		Erigeron canadensis	A 158	0.6	Onobrychis viciifolia	E	16 (43)
		Crassula lycopodioides	A 119	0.5	Robinia pseudoacacia	A	12 (43)
	Mesic	Erigeron annuus	A 819	2.5	Erigeron canadensis	A	22 (45)
	(32740) Trifolium hybridum	C 533	1.6	Erigeron annuus	A	18 (45)
		Medicago sativa	C 435	1.3	Matricaria discoidea	A	18 (45)
		Erigeron canadensis	A 366	1.1	Medicago sativa	C	18 (45)
		Onobrychis viciifolia	E 320	1.0	Juncus tenuis	A	16 (45)
	Wet	Solidago gigantea	A 417	1.8	Bidens frondosus	A	16 (47)
	(22763) Epilobium ciliatum	A 324	1.4	Erigeron annuus	A	16 (47)
		Impatiens glandulifera	A 283	1.2	Erigeron canadensis	A	16 (47)
		Trifolium hybridum	C 254	1.1	Epilobium ciliatum	A	15 (47)
		Bidens frondosus	A 188	0.8	Impatiens glandulifera	A	15 (47)
	Alpine	Onobrychis viciifolia	E 9	0.1			1 (26)
	(6086)	Amorpha fruticosa	A 4	0.1			1 (26)
		Alchemilla conjuncta	E 3	0.0			1 (26)
		Cota triumfettii	E 2	0.0			1 (26)
		Oenothera glazioviana	A 2	0.0			1 (26)
	Saline	Grindelia squarrosa	A 17	1.7	Erigeron canadensis	A	6 (22)
	(988)	Xanthium orientale	A 17	1.7	Symphyotrichum squamatum A	3 (22)
		Erigeron canadensis	A 8	0.8	Xanthium orientale	A	3 (22)
		Ambrosia artemisiifolia	A 7	0.7	Xanthium spinosum	A	3 (22)
		Symphyotrichum squamatum A 4	0.4	Cotula coronopifolia	A	2 (22)

  Neophyte invasions in European grasslands. Journal of Vegetation Science. Overview of contributing vegetation-plot databases, number (n) and percentage (%) of plots used in the final dataset(N = 97,417). Databases are 20 arranged according to the codes in the GIVD database. Neophyte invasions in European grasslands. Journal of Vegetation Science. Overview of the classification of vegetation plots into habitat types according to the European Nature Information System (EUNIS) and their 26 assignment into broad vegetation types. Habitat group combines vegetation types of similar ecology and vegetation characteristics. The total number of plots in 27 the final dataset was 97,417. Asterisks indicate grassland plots from Denmark (National Vegetation Database of Denmark) that were not classified to EUNIS habitats 28 but assigned into broad vegetation types based on their similarity in species composition (Bray-Curtis index) to plots from other countries (see Methods for details). Neophyte invasions in European grasslands. Journal of Vegetation Science. List of regions. Regions marked by an asterisk have less than 100 plots; therefore, statistics for these regions are less representative. The European 33 part of the Russian Federation is divided into regions according to the Euro+Med PlantBase. Neophyte invasions in European grasslands. Journal of Vegetation Science. Overview of the species-pool data and relative frequency of invaded plots (plots with the presence of at least one neophyte) for Europe and individual 252 countries. For Europe, we included into native category only species considered native in all regions, while species of European origin considered as a neophyte in 253 at least one country were included in the neophyte numbers. Regions marked by an asterisk have less than 100 plots; therefore, statistics for these regions are 254 less representative. The European part of the Russian Federation is divided into regions according to the Euro+Med PlantBase. Neophyte in European grasslands. Journal of Vegetation Science. Overview of the species-pool data and relative frequency of invaded plots (plots with the presence of at least one neophyte) for habitat groups and 268 EUNIS habitats. Note that number of all species refers to taxa at the species level. EUNIS habitats marked by an asterisk have less than 100 plots; therefore, 269 statistics for these habitats are less representative. Non-classified plots from Denmark are not included in the EUNIS habitat statistics (in total 1885 plots). Neophyte invasions in European grasslands. Journal of Vegetation Science. List of neophyte species recorded in individual regions. Species are listed alphabetically and their frequency in regional subsets is shown under No. 283 of plots in which the species was recorded. Subspecies were listed only if recorded in the vegetation plot and considered as neophytes for the given region. Russia 284 refers to European part of Russia, which is in our study subdivided into seven regions. All the regions are delimited following the Euro+Med PlantBase. Note that 285 some regions are considered separately due to their specific history or ecology (e.g. Crete). Regions with a low number of plots (less than 100) are marked by an 286 asterisk. Origin 'A' refers to non-European origin, 'E' to European origin and 'C' to other origin (anecophytes, hybrids).

	17 18 Axmanová, I. et al. 19 Supporting information to the paper Appendix S1. GIVD database name Database of non-forest vegetation of the Southern Urals Nenets Tundra Nordic-Baltic Grassland Vegetation Database (NBGVD) Iberian and Macaronesian Vegetation Information System (SIVIM) Iberian and Macaronesian Vegetation Information System (SIVIM) -Alpine grasslands Iberian and Macaronesian Vegetation Information System (SIVIM) -Catalonia Iberian and Macaronesian Vegetation Information System (SIVIM) -Grasslands Vegetation-Plot Database of the University of the Basque Country (BIOVEG) Balkan Dry Grasslands Database Mediterranean Ammophiletea database European Coastal Vegetation Database The Nordic Vegetation Database Balkan Vegetation Database European Mire Vegetation Database High Mediterranean Mountains Database Masaryk University's Gap-Filling Database of European Vegetation Vegetation Database of Albania Austrian Vegetation Database INBOVEG Bulgarian Vegetation Database Monitoring Effectiveness of Habitat Conservation in Switzerland F o r R e v i e w GIVD code n 00-RU-006 1702 1.7 Sergey Yamalov % Custodian AS-RU-005 9 <0.1 Igor Lavrinenko EU-00-002 1392 1.4 Jürgen Dengler EU-00-004 1 <0.1 Xavier Font EU-00-004 274 0.3 Borja Jiménez-Alfaro EU-00-004 55 0.1 Xavier Font EU-00-004 1549 1.6 Maria Pilar Rodríguez-Rojo EU-00-011 2225 2.3 Idoia Biurrun EU-00-013 2577 2.6 Kiril Vassilev EU-00-016 455 0.5 Corrado Marcenò EU-00-017 61 0.1 John Janssen EU-00-018 806 0.8 Jonathan Lenoir EU-00-019 1008 1.0 Kiril Vassilev EU-00-022 23 <0.1 Tomáš Peterka EU-00-029 132 0.1 Gianpietro Giusso del Galdo Corrado Marcenò Deputy custodian Mariya Lebedeva Łukasz Kozub Xavier Font Xavier Font Itziar García-Mijangos Armin Macanović Borja Jiménez-Alfaro Jens-Christian Svenning Hristo Pedashenko Martin Jiroušek EU-00-031 1 <0.1 Milan Chytrý Ilona Knollová EU-AL-001 29 <0.1 Michele De Sanctis Giuliano Fanelli EU-AT-001 4739 4.9 Wolfgang Willner EU-BE-002 117 0.1 Els De Bie EU-BG-001 615 0.6 Iva Apostolova Desislava Sopotlieva EU-CH-011 1807 1.9 Ariel Bergamini Steffen Boch O n l y Czech National Phytosociological Database EU-CZ-001 5324 5.5 Milan Chytrý Ilona Knollová VegMV EU-DE-001 3806 3.9 Florian Jansen Christian Berg VegetWeb Germany 1 EU-DE-013 1848 1.9 Florian Jansen Jörg Ewald VegetWeb Germany 2 (Tüxen's archive) EU-DE-013 31 <0.1 Friedemann Goral Florian Jansen German Vegetation Reference Database (GVRD) EU-DE-014 5965 6.1 Ute Jandt Helge Bruelheide German Grassland Vegetation Database (GrassVeg.DE) EU-DE-020 1646 1.7 Jürgen Dengler Ricarda Pätsch Database Schleswig-Holstein (Northern Germany) EU-DE-040 332 0.3 Joachim Schrautzer National Vegetation Database of Denmark EU-DK-002 1885 1.9 Jesper Erenskjold Moeslund Rasmus Ejrnaes Iberian and Macaronesian Vegetation Information System (SIVIM) -Wetlands EU-ES-001 98 0.1 Aaron Pérez-Haase Xavier Font SOPHY EU-FR-003 20404 20.9 Henry Brisse Patrice de Ruffray UK National Vegetation Classification Database EU-GB-001 2016 2.1 John S. Rodwell KRITI EU-GR-001 93 0.1 Erwin Bergmeier Hellenic Natura 2000 Vegetation Database (HelNatVeg) EU-GR-005 126 0.1 Panayotis Dimopoulos Ioannis Tsiripidis Croatian Vegetation Database EU-HR-002 513 0.5 Željko Škvorc Daniel Krstonošić CoenoDat Hungarian Phytosociological Database EU-HU-003 236 0.2 János Csiky Zoltán Botta-Dukát Irish Vegetation Database EU-IE-001 143 0.1 Úna FitzPatrick Lynda Weekes VegItaly EU-IT-001 483 0.5 Roberto Venanzoni Flavia Landucci Vegetation database of Habitats in the Italian Alps -HabItAlp EU-IT-010 292 0.3 Laura Casella Pierangela Angelini Vegetation Plot Database -Sapienza University of Rome EU-IT-011 1572 1.6 Emiliano Agrillo Fabio Attorre VIOLA EU-IT-019 416 0.4 Angela Stanisci Maria Laura Carranza Lithuanian vegetation Database EU-LT-001 462 0.5 Valerius Rašomavičius Domas Uogintas Semi-natural Grassland Vegetation Database of Latvia EU-LV-001 882 0.9 Solvita Rūsiņa Vegetation Database of the Republic of Macedonia EU-MK-001 29 <0.1 Renata Ćušterevska Dutch National Vegetation Database EU-NL-001 1617 1.7 Stephan Hennekens Joop Schaminée Dutch Military Ranges Vegetation Database (DUMIRA) EU-NL-003 71 0.1 Iris de Ronde Rense Haveman Polish Vegetation Database EU-PL-001 10433 10.7 Zygmunt Kącki F o r R e v i e w O n l y Romanian Grassland Database EU-RO-008 5118 5.3 Eszter Ruprecht Kiril Vassilev Vegetation Database Grassland Vegetation of Serbia EU-RS-002 1545 1.6 Svetlana Aćić Zora Dajić Stevanović Database of Forest Vegetation in Republic of Serbia EU-RS-003 34 <0.1 Mirjana Krstivojević Ćuk & Vegetation Database of Northern Part of Serbia (AP Vojvodina) EU-RS-004 Lower Volga Valley Phytosociological Database EU-RU-002 684 0.7 Valentin Golub Andrei Chuvashov Vegetation Database of the Volga and the Ural Rivers Basins EU-RU-003 299 0.3 Tatiana Lysenko Vegetation Database of Tatarstan EU-RU-011 485 0.5 Vadim Prokhorov Vegetation Database of Slovenia EU-SI-001 1256 1.3 Urban Šilc Filip Kuzmič Slovak Vegetation Database EU-SK-001 3388 3.5 Milan Valachovič Jozef Šibík Ukrainian Grassland Database EU-UA-001 2331 2.4 Anna Kuzemko Yulia Vashenyak Halophytic and coastal vegetation database of Ukraine EU-UA-005 250 0.3 Tetiana Dziuba Dmytro Dubyna Vegetation Database of Ukraine and Adjacent Parts of Russia EU-UA-006 1018 1.0 Viktor Onyshchenko Vitaliy Kolomiychuk Steppe Vegetation Rostov Region Database -567 0.6 Olga Demina Teberda -Caucasus Database -142 0.1 Vladimir Onipchenko Alexei Egorov 21 22 F o r R e v i e w O n l y 23 Supporting information to the paper 24 Axmanová, I. et al. 25 Appendix S2. EUNIS code EUNIS habitat Habitat group Plot number N15 Atlantic and Baltic coastal dune grassland (grey dune) Sandy 468 N16 Mediterranean and Macaronesian coastal dune grassland (grey dune) Sandy 714 N17 Black Sea coastal dune grassland (grey dune) Sandy 390 R11 Pannonian and Pontic sandy steppe Sandy 192 R12 Cryptogam-and annual-dominated vegetation on siliceous rock outcrops Rocky 235 R13 Cryptogam-and annual-dominated vegetation on calcareous and ultramafic rock outcrops Rocky 687 R14 Perennial rocky grassland of the Italian Peninsula Rocky 371 R15 Continental dry rocky steppic grassland and dwarf scrub on chalk outcrops Rocky 123 R16 Perennial rocky grassland of Central and South-Eastern Europe Rocky 1615 R17 Heavy-metal dry grassland of the Balkans Rocky 21 R18 Perennial rocky calcareous grassland of subatlantic-submediterranean Europe Rocky 1610 R19 Dry steppic submediterranean pasture of the Amphi-Adriatic region Dry 223 R1A Semi-dry perennial calcareous grassland (meadow steppe) Dry 16100 R1B Continental dry grassland (true steppe) Dry 6176 R1C Desert steppe Dry 440 R1D Mediterranean closely grazed dry grassland Dry 149 R1E Mediterranean tall perennial dry grassland Dry 777 R1F Mediterranean annual-rich dry grassland Dry 70 R1G Iberian oromediterranean siliceous dry grassland Oromediterranean 65 R1H Iberian oromediterranean basiphilous dry grassland Oromediterranean 209 R1J Cyrno-Sardean oromediterranean siliceous dry grassland Oromediterranean 28 R1K Balkan and Anatolian oromediterranean dry grassland Oromediterranean 78 R1M Lowland to montane, dry to mesic grassland usually dominated by Nardus stricta Mesic 826 R1N Open Iberian supramediterranean dry acid and neutral grassland Dry 15 R1P Oceanic to subcontinental inland sand grassland on dry acid and neutral soils Sandy 1989 R1Q Inland sanddrift and dune with siliceous grassland Sandy 606 F o r R e v i e w O n l y R1R Mediterranean to Atlantic open, dry, acid and neutral grassland Dry 561 R1S Heavy-metal grassland in Western and Central Europe Rocky 9 R21 Mesic permanent pasture of lowlands and mountains Mesic 6542 R22 Low and medium altitude hay meadow Mesic 22885 R23 Mountain hay meadow Mesic 1400 R24 Iberian summer pasture (vallicar) Dry 15 R31 Mediterranean tall humid inland grassland Wet 313 R32 Mediterranean short moist grassland of lowlands Wet 14 R33 Mediterranean short moist grassland of mountains Mesic 147 R34 Submediterranean moist meadow Wet 244 R35 Moist or wet mesotrophic to eutrophic hay meadow Wet 9408 R36 Moist or wet mesotrophic to eutrophic pasture Wet 3460 R37 Temperate and boreal moist or wet oligotrophic grassland Wet 2946 R41 Snow-bed vegetation Alpine 511 R42 Boreal and arctic acidophilous alpine grassland Alpine 131 R43 Temperate acidophilous alpine grassland Alpine 3157 R44 Arctic-alpine calcareous grassland Alpine 1585 R45 Alpine and subalpine calcareous grassland of the Balkans and Apennines Alpine 702 R51 Thermophilous forest fringe of base-rich soils Dry 477 R52 Forest fringe of acidic nutrient-poor soils Dry 305 R55 Lowland moist or wet tall-herb and fern fringe Wet 4723 R56 Montane to subalpine moist or wet tall-herb and fern fringe Wet 832 R61 Mediterranean inland salt steppe Saline 79 R62 Continental inland salt steppe Saline 460 R63 Temperate inland salt marsh Saline 357 R64 Semi-desert salt pan Saline 81 R65 Continental subsaline alluvial pasture and meadow Saline 11 * NA Dry 122 * NA Mesic 940 * NA Wet 823 29 F o r R e v i e w O n l y 30 Supporting information to the paper 31 Axmanová, I. et al. 32 Appendix S3. 34 Region Number of plots Region Number of plots Region Number of plots Albania* 29 Hungary 273 Russia (Kaliningrad)* 3 Austria 4767 Ireland 147 Russia (North) 19 Baleares* 7 Italy 2691 Russia (Northwest) 44 Belarus* 42 Kosovo 259 Russia (South) 1294 Belgium 137 Latvia 912 Romania 5153 Bosnia and Herzegovina 243 Lithuania 468 Sardinia 162 Bulgaria 3466 Luxembourg* 12 Serbia 1531 Corsica 268 Macedonia* 71 Sicily and Malta 224 Crete* 98 Moldova 201 Slovakia 3366 Croatia 537 Montenegro* 42 Slovenia 1267 Czech Republic 5215 Netherlands 1682 Spain 4361 Denmark 1885 Norway 348 Sweden 390 Estonia 955 Poland 10490 Switzerland 2376 Finland 430 Portugal 160 Ukraine 3276 France 19069 Russia (Central)* 42 United Kingdom 2019 Germany 13882 Russia (Caucasus) 289 Greece 480 Russia (East) 2335 35 36 37 1 2 3 4 5 6 7 8 1 2 3 4 5 6 7 8 1 2 3 4 5 6 7 8 1 2 3 4 5 6 7 8 1 2 3 4 5 6 7 8 1 2 3 4 5 6 7 8 F o r R e v i e w O n l y 249 Supporting information to the paper 250 Axmanová, I. et al. 251 Appendix S6. Region Number of all species Number of native species and archaeophytes Number of neophytes Non-European origin European origin Other (anecophytes, hybrids) Relative frequency of neophytes in the species pool [%] Relative frequency of non-European neophytes [%] All plots Plots with native taxa only (no neophytes) Plots with at least one neophyte present (invaded plots) % of invaded plots % of plots invaded by non-European neophytes EUROPE 8212 7676 536 249 251 36 6.5 3.0 97417 85871 11546 11.9 7.1 Albania* 247 244 3 2 1 0 1.2 0.8 29 26 3 10.3 6.9 Austria 1870 1806 64 29 27 8 3.4 1.6 4767 3877 890 18.7 9.8 Baleares* 20 20 0 0 0 0 0.0 0.0 7 7 0 0.0 0.0 Belarus* 229 218 11 6 2 3 4.8 2.6 42 21 21 50.0 35.7 Belgium 372 355 17 9 7 1 4.6 2.4 137 106 31 22.6 19.7 Bosnia and Herzegovina 870 860 10 8 1 1 1.1 0.9 243 230 13 5.3 4.1 Bulgaria 2215 2194 21 17 1 3 0.9 0.8 3466 3368 98 2.8 2.6 Corsica 715 705 10 9 1 0 1.4 1.3 268 246 22 8.2 7.8 Crete* 639 628 11 10 0 1 1.7 1.6 98 92 6 6.1 5.1 Croatia 1061 1035 26 23 2 1 2.5 2.2 537 334 203 37.8 35.2 Czech Republic 1570 1489 81 44 30 7 5.2 2.8 5215 4492 723 13.9 7.4 Denmark 783 713 70 23 40 7 8.9 2.9 1885 1475 410 21.8 12.1 Estonia 377 371 6 0 5 1 1.6 0.0 955 943 12 1.3 0.0 Finland 433 413 20 3 16 1 4.6 0.7 430 159 271 63.0 11.4 France 3171 3050 121 86 26 9 3.8 2.7 19069 18028 1041 5.5 3.8 Germany 1568 1458 110 47 56 7 7.0 3.0 13882 12111 1771 12.8 5.6 F o r R e v i e w O n l y Greece 1228 1222 6 6 0 0 0.5 0.5 480 466 14 2.9 2.9 Hungary 789 775 14 10 3 1 1.8 1.3 273 228 45 16.5 15.8 Ireland 344 338 6 4 2 0 1.7 1.2 147 135 12 8.2 6.1 Italy 2471 2421 50 44 6 0 2.0 1.8 2691 2312 379 14.1 8.2 Kosovo 622 621 1 0 0 1 0.2 0.0 259 257 2 0.8 0.0 Latvia 557 528 29 14 12 3 5.2 2.5 912 813 99 10.9 5.2 Lithuania 602 568 34 21 11 2 5.6 3.5 468 353 115 24.6 19.2 Luxembourg* 102 102 0 0 0 0 0.0 0.0 12 12 0 0.0 0.0 Macedonia* 590 589 1 0 1 0 0.2 0.0 71 66 5 7.0 0.0 Moldova 489 475 14 10 3 1 2.9 2.0 201 166 35 17.4 15.9 Montenegro* 283 283 0 0 0 0 0.0 0.0 42 42 0 0.0 0.0 Netherlands 686 641 45 26 14 5 6.6 3.8 1682 1504 178 10.6 8.1 Norway 342 338 4 0 4 0 1.2 0.0 348 341 7 2.0 0.0 Poland 1589 1471 118 62 47 9 7.4 3.9 10490 8646 1844 17.6 12.3 Portugal 506 500 6 6 0 0 1.2 1.2 160 127 33 20.6 20.6 Russia (Central)* 331 329 2 0 2 0 0.6 0.0 42 33 9 21.4 0.0 Russia (Caucasus) 762 752 10 8 2 0 1.3 1.0 289 276 13 4.5 4.2 Russia (East) 1271 1244 27 18 6 3 2.1 1.4 2335 2210 125 5.4 3.3 Russia (Kaliningrad)* 43 40 3 3 0 0 7.0 7.0 3 0 3 100.0 100.0 Russia (North) 118 118 0 0 0 0 0.0 0.0 19 19 0 0.0 0.0 Russia (Northwest) 165 163 2 1 1 0 1.2 0.6 44 41 3 6.8 2.3 Russia (South) 898 876 22 16 4 2 2.4 1.8 1294 1161 133 10.3 3.6 Romania 2125 2073 52 42 8 2 2.4 2.0 5153 4121 1032 20.0 9.4 Sardinia 475 466 9 8 0 1 1.9 1.7 162 152 10 6.2 5.6 Serbia 1416 1403 13 13 0 0 0.9 0.9 1531 1462 69 4.5 4.5 Sicily and Malta 660 648 12 11 1 0 1.8 1.7 224 197 27 12.1 11.6 Slovakia 1686 1637 49 25 20 4 2.9 1.5 3366 3096 270 8.0 3.7 Slovenia 1410 1374 36 30 3 3 2.6 2.1 1267 971 296 23.4 20.9 Spain 2747 2671 76 61 8 7 2.8 2.2 4361 4022 339 7.8 6.6 Sweden 401 391 10 1 8 1 2.5 0.2 390 368 22 5.6 0.3 Switzerland 1521 1482 39 22 15 2 2.6 1.4 2376 2129 247 10.4 4.6 Ukraine 1838 1767 71 42 22 7 3.9 2.3 3276 2713 563 17.2 11.8 United Kingdom 773 729 44 13 26 5 5.7 1.7 2019 1917 102 5.1 2.3 255 256 F o r R e v i e w O n l y F o r R e v i e w O n l y 265 Supporting information to the paper 266 Axmanová, I. et al. 267 Appendix S8. Habitat Number of all species Number of native species Number of neophytes Non-European origin European origin Other (anecophytes, hybrids) Relative frequency of all neophytes in the species pool [%] Relative frequency of non-European neophytes [%] All plots Plots with only native taxa (no neophytes) Plots with at least one neophyte present (invaded plots) % of invaded plots % of plots invaded by non-European Habitat group Sandy 2150 2020 130 80 46 4 6.1 3.7 4359 3237 1122 25.7 21.8 neophytes Rocky 3354 3293 61 24 33 4 1.8 0.7 4671 4408 263 5.6 2.3 Dry 6016 5798 218 89 111 18 3.7 1.5 25430 22701 2729 10.7 3.9 Oromediterranean 957 957 0 0 0 0 0.0 0.0 380 380 0 0.0 0.0 Mesic 4014 3736 278 115 141 22 7.2 2.9 32740 28327 4413 13.5 7.5 Wet 3584 3323 261 158 85 18 7.4 4.4 22763 19847 2916 12.8 10.2 Alpine 3180 3167 13 6 7 0 0.4 0.2 6086 6056 30 0.5 0.2 Saline 846 828 18 13 2 3 2.1 1.5 988 915 73 7.4 6.4 F o r R e v i e w O n l y N15 Atlantic and Baltic coastal dune grassland (grey dune) 470 439 31 19 12 0 6.6 4.0 468 393 75 16.0 13.7 N16 Mediterranean and Macaronesian coastal dune grassland (grey dune) 692 658 34 32 2 0 4.9 4.6 714 552 162 22.7 22.5 N17 Black Sea coastal dune grassland (grey dune) 463 446 17 12 5 0 3.7 2.6 390 305 85 21.8 19.0 R11 Pannonian and Pontic sandy steppe 534 516 18 15 2 1 3.4 2.8 192 153 39 20.3 19.3 R12 Cryptogam-and annual-dominated vegetation on siliceous rock outcrops 628 619 9 6 3 0 1.4 1.0 235 192 43 18.3 11.9 R13 Cryptogam-and annual-dominated vegetation on calcareous and ultramafic rock outcrops 1245 1213 32 17 14 1 2.6 1.4 687 623 64 9.3 5.7 R14 Perennial rocky grassland of the Italian Peninsula 874 872 2 0 2 0 0.2 0.0 371 353 18 4.9 0.0 R15 Continental dry rocky steppic grassland and dwarf scrub on chalk outcrops 443 440 3 1 2 0 0.7 0.2 123 116 7 5.7 3.3 R16 Perennial rocky grassland of Central and South-Eastern Europe 1969 1946 23 6 15 2 1.2 0.3 1615 1541 74 4.6 1.5 R17* Heavy-metal dry grassland of the Balkans 243 243 0 0 0 0 0.0 0.0 21 21 0 0.0 0.0 R18 Perennial rocky calcareous grassland of subatlantic-submediterranean Europe 1483 1465 18 8 8 2 1.2 0.5 1610 1553 57 3.5 0.8 R19 Dry steppic submediterranean pasture of the Amphi-Adriatic region 995 993 2 0 2 0 0.2 0.0 223 218 5 2.2 0.0 R1A Semi-dry perennial calcareous grassland (meadow steppe) 4145 4003 142 55 74 13 3.4 1.3 16100 14262 1838 11.4 3.1 R1B Continental dry grassland (true steppe) 2624 2545 79 36 38 5 3.0 1.4 6176 5598 578 9.4 4.7 R1C Desert steppe 533 528 5 2 2 1 0.9 0.4 440 434 6 1.4 F o r R e v i e w O n l y R1E Mediterranean tall perennial dry grassland 1668 1647 21 13 6 2 1.3 0.8 777 727 50 6.4 3.3 R1F* Mediterranean annual-rich dry grassland 455 452 3 2 1 0 0.7 0.4 70 61 9 12.9 7.1 R1G* Iberian oromediterranean siliceous dry grassland 175 175 0 0 0 0 0.0 0.0 65 65 0 0.0 0.0 R1H Iberian oromediterranean basiphilous dry grassland 604 604 0 0 0 0 0.0 0.0 209 209 0 0.0 0.0 R1J* Cyrno-Sardean oromediterranean siliceous dry grassland 99 99 0 0 0 0 0.0 0.0 28 28 0 0.0 0.0 R1K* Balkan and Anatolian oromediterranean dry grassland 239 239 0 0 0 0 0.0 0.0 78 78 0 0.0 0.0 R1M Lowland to montane, dry to mesic grassland usually dominated by Nardus stricta 889 877 12 9 3 0 1.3 1.0 826 807 19 2.3 1.9 R1N* Open Iberian supramediterranean dry acid and neutral grassland 182 182 0 0 0 0 0.0 0.0 15 15 0 0.0 0.0 R1P Oceanic to subcontinental inland sand grassland on dry acid and neutral soils 1219 1152 67 33 30 4 5.5 2.7 1989 1371 618 31.1 24.8 R1Q Inland sanddrift and dune with siliceous grassland 397 376 21 10 11 0 5.3 2.5 606 463 143 23.6 19.6 R1R Mediterranean to Atlantic open, dry, acid and neutral grassland 968 959 9 6 3 0 0.9 0.6 561 445 116 20.7 17.3 R1S* F o r R e v i e w O n l y R31 Mediterranean tall humid inland grassland 695 669 26 24 1 1 3.7 3.5 313 249 64 20.4 19.5 R32* Mediterranean short moist grassland of lowlands 187 186 1 0 0 1 0.5 0.0 14 13 1 7.1 0.0 R33 Mediterranean short moist grassland of mountains 313 313 0 0 0 0 0.0 0.0 147 147 0 0.0 0.0 R34 Submediterranean moist meadow 575 569 6 3 1 2 1.0 0.5 244 231 13 5.3 3.7 R35 Moist or wet mesotrophic to eutrophic hay meadow 1849 1754 95 57 32 6 5.1 3.1 9408 8521 887 9.4 5.6 R36 Moist or wet mesotrophic to eutrophic pasture 1381 1288 93 69 15 9 6.7 5.0 3460 2894 566 16.4 14.5 R37 Temperate and boreal moist or wet oligotrophic grassland 1425 1390 35 19 15 1 2.5 1.3 2946 2815 131 4.4 3.5 R41 Snow-bed vegetation 791 790 1 0 1 0 0.1 0.0 511 510 1 0.2 0.0 R42 Boreal and arctic acidophilous alpine grassland 199 198 1 0 1 0 0.5 0.0 131 130 1 0.8 0.0 R43 Temperate acidophilous alpine grassland 1899 1892 7 5 2 0 0.4 0.3 3157 3146 11 0.3 0.3 R44 Arctic-alpine calcareous grassland 1837 1833 4 1 3 0 0.2 0.1 1585 1578 7 0.4 0.1 R45 Alpine and subalpine calcareous grassland of the Balkans and Apennines 1443 1441 2 0 2 0 0.1 0.0 702 692 10 1.4 0.0 R51 Thermophilous forest fringe of base-rich soils 1278 1256 22 10 9 3 1.7 0.8 477 439 38 8.0 4.2 F o r R e v i e w O n l y R63 Temperate inland salt marsh 475 461 14 10 2 2 2.9 2.1 357 327 30 8.4 6.2 R64* Semi-desert salt pan 121 119 2 2 0 0 1.7 1.7 81 79 2 2.5 2.5 R65* Continental subsaline alluvial pasture and meadow 53 50 3 2 1 0 5.7 3.8 11 9 2 18.2 18.2 270 F o r R e v i e w O n l y F o r R e v i e w O n l y 280 Supporting information to the paper 281 Region Taxon Origin No. of plots % of plots Subsp. 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Axmanová, I. et al. 282 Austria Chenopodium hircinum A 1 0.02 Austria Iberis pinnata E 4 0.08 Austria Impatiens glandulifera A 35 0.73 Austria Impatiens parviflora A 25 0.52 Austria Juncus tenuis A 7 0.15 Austria Ligularia dentata A 1 0.02 Austria Symphyotrichum novi-belgii A 3 0.06 Austria Symphyotrichum salignum A 1 0.02 Austria Syringa vulgaris E 5 0.10 Austria Veronica filiformis E 1 0.02 Austria Veronica peregrina A 1 0.02 Austria Veronica persica A 9 0.19 Bosnia and Herzegovina Amaranthus retroflexus A 1 0.41 Bosnia and Herzegovina Ambrosia artemisiifolia A 2 0.82 Bosnia and Herzegovina Bolboschoenus fluviatilis A 1 0.41 Bosnia and Herzegovina Centaurea cariensis E 1 0.41 Bosnia and Herzegovina Erigeron annuus A 7 2.88 Bosnia and Herzegovina Impatiens glandulifera A 1 0.41 Corsica Erigeron bonariensis A 1 0.37 Corsica Erigeron sumatrensis A 1 0.37 Corsica Paspalum distichum A 4 1.49 Corsica Pinus pinea E 1 0.37 Corsica Symphyotrichum squamatum A 6 2.24 Corsica Xanthium orientale A 2 0.75 Croatia Panicum capillare A 1 0.19 Croatia Potentilla intermedia E 1 0.19 Croatia Rudbeckia laciniata A 3 0.56 Croatia Solidago canadensis A 5 0.93 Croatia Solidago gigantea A 11 2.05 Croatia Sorghum halepense A 2 0.37 Czech Republic Geranium pyrenaicum E 2 0.04 Czech Republic Helianthus tuberosus A 8 0.15 Czech Republic Heracleum mantegazzianum E 2 0.04 Czech Republic Hesperis matronalis E 2 0.04 Czech Republic Chenopodium strictum E 2 0.04 Czech Republic Impatiens glandulifera A 62 1.19 Czech Republic Reynoutria x bohemica A 2 0.04 Czech Republic Robinia pseudoacacia A 13 0.25 Czech Republic Rudbeckia laciniata A 1 0.02 Czech Republic Rumex thyrsiflorus E 41 0.79 Czech Republic Sedum spurium E 1 0.02 Czech Republic Senecio leucanthemifolius E 1 0.02 Denmark Epilobium ciliatum A 146 7.75 subsp. adenocaulon, subsp. ciliatum, subsp. glandulosum Denmark Epilobium komarovianum A 3 0.16 Denmark Erigeron canadensis A 4 0.21 Denmark Falcaria vulgaris E 1 0.05 Denmark Heracleum mantegazzianum E 15 0.80 Denmark Heracleum sphondylium E 3 0.16 Denmark Ribes uva-crispa E 6 0.32 Denmark Rosa rugosa A 27 1.43 Denmark Rubus laciniatus C 1 0.05 Denmark Rubus ulmifolius E 1 0.05 Denmark Rudbeckia A 1 0.05 Denmark Salix alba E 1 0.05 Finland Holcus mollis E 2 0.47 Finland Chaerophyllum prescottii E 8 1.86 Finland Lolium multiflorum E 2 0.47 Finland Lolium perenne E 15 3.49 Finland Lupinus polyphyllus A 1 0.23 Finland Matricaria discoidea A 19 4.42 France Bidens connatus A 1 0.01 France Bidens frondosus A 31 0.16 France Brassica elongata E 1 0.01 France Brassica napus C 2 0.01 France Bromus brachystachys E 2 0.01 France Buddleja davidii A 3 0.02 France Impatiens glandulifera A 41 0.22 France Impatiens parviflora A 5 0.03 France Inula helenium E 8 0.04 France Juglans regia E 38 0.20 France Juncus tenuis A 46 0.24 France Lathyrus hirsutus E 25 0.13 France Prunus dulcis C 3 0.02 France Prunus serotina A 7 0.04 France Pseudotsuga menziesii A 6 0.03 France Quercus rubra A 5 0.03 France Reynoutria japonica A 8 0.04 France Ricinus communis A 1 0.01 Germany Agrostis castellana E 1 0.01 Germany Alchemilla mollis E 20 0.14 Germany Amaranthus retroflexus A 2 0.01 Germany Ambrosia artemisiifolia A 3 0.02 Germany Anthoxanthum aristatum E 18 0.13 Germany Armoracia rusticana E 4 0.03 Germany Helianthus annuus A 2 0.01 Germany Helianthus tuberosus A 1 0.01 Germany Helminthotheca echioides E 1 0.01 Germany Heracleum mantegazzianum E 1 0.01 Germany Hesperis matronalis E 5 0.04 Germany Chenopodium strictum E 1 0.01 Germany Prunus domestica C 22 0.16 Germany Prunus serotina A 7 0.05 Germany Pseudotsuga menziesii A 1 0.01 Germany Quercus rubra A 3 0.02 Germany Reynoutria sachalinensis A 1 0.01 Germany Robinia pseudoacacia A 12 0.09 Germany Vicia grandiflora E 1 0.01 Germany Vicia pannonica E 2 0.01 subsp. pannonica Germany Vicia villosa E 41 0.30 subsp. villosa Germany Xanthium orientale A 4 0.03 Germany Zea mays A 1 0.01 Greece Cenchrus spinifex A 1 0.21 Italy Ambrosia tenuifolia A 1 0.04 Italy Amorpha sp. A 4 0.15 Italy Amorpha fruticosa A 10 0.37 Italy Artemisia verlotiorum A 9 0.33 Italy Bidens frondosus A 12 0.45 Italy Carpobrotus acinaciformis A 2 0.07 Italy Phalaris canariensis E 2 0.07 Italy Phytolacca americana A 1 0.04 Italy Potentilla norvegica E 2 0.07 Italy Reynoutria japonica A 1 0.04 Italy Robinia pseudoacacia A 2 0.07 Italy Senecio inaequidens A 1 0.04 Latvia Potentilla inclinata E 9 0.99 Latvia Prunus avium E 1 0.11 Latvia Rumex confertus E 7 0.77 Latvia Saponaria officinalis E 4 0.44 Latvia Senecio leucanthemifolius E 6 0.66 Latvia Solidago canadensis A 6 0.66 Lithuania Rosa rugosa A 3 0.64 Lithuania Rudbeckia sp. A 1 0.21 Lithuania Salix euxina E 1 0.21 Lithuania Saponaria officinalis E 7 1.50 Lithuania Sedum album E 1 0.21 Lithuania Sedum rupestre E 1 0.21 Netherlands Elodea canadensis A 2 0.12 Netherlands Elodea nuttallii A 3 0.18 Netherlands Epilobium ciliatum A 8 0.48 Netherlands Erigeron canadensis A 26 1.55 Netherlands Galinsoga quadriradiata A 1 0.06 Netherlands Geranium pyrenaicum E 1 0.06 Norway Achillea ptarmica E 1 0.29 Norway Noccaea caerulescens E 1 0.29 Norway Potentilla anglica E 1 0.29 Norway Potentilla reptans E 4 1.15 Poland Acer negundo A 3 0.03 Poland Acer saccharinum A 2 0.02 Poland Euonymus fortunei A 1 0.01 Poland Euthamia graminifolia A 7 0.07 Poland Forsythia suspensa A 1 0.01 Poland Fragaria ananassa C 1 0.01 Poland Fraxinus pennsylvanica A 4 0.04 Poland Galinsoga parviflora A 18 0.17 Poland Narcissus poeticus E 1 0.01 Poland Oenothera sp. A 1 0.01 Poland Oenothera ammophila A 1 0.01 Poland Oenothera biennis A 62 0.59 Poland Oenothera parviflora A 1 0.01 Poland Oenothera rubricaulis A 38 0.36 Poland Solidago canadensis A 112 1.07 Poland Solidago gigantea A 226 2.15 Poland Spiraea japonica A 1 0.01 Poland Spiraea tomentosa A 1 0.01 Poland Symphoricarpos albus A 1 0.01 Poland Symphyotrichum lanceolatum A 8 0.08 Russia (Caucasus) Veronica peregrina A 1 0.35 Russia (Caucasus) Xanthium orientale A 3 1.04 Russia (Caucasus) Xanthium pungens A 2 0.69 Russia (East) Acer negundo A 8 0.34 Russia (East) Aconitum nemorum A 2 0.09 Russia (East) Caragana arborescens A 1 0.04 Russia (NorthWest) Schedonorus arundinaceus E 2 4.55 Russia (South) Acorus calamus A 1 0.08 Russia (South) Amaranthus albus A 6 0.46 Russia (South) Amaranthus retroflexus A 5 0.39 Russia (South) Ambrosia artemisiifolia A 6 0.46 Russia (South) Amorpha fruticosa A 2 0.15 Sicily and Malta Erigeron canadensis A 3 1.34 Sicily and Malta Megathyrsus maximus A 4 1.79 Sicily and Malta Opuntia ficus-indica A 2 0.89 Sicily and Malta Oxalis pes-caprae A 9 4.02 Sicily and Malta Saccharum spontaneum A 6 2.68 Sicily and Malta Symphyotrichum squamatum A 2 0.89 Slovakia Oenothera biennis A 1 0.03 Slovakia Oenothera villosa A 2 0.06 Slovakia Onobrychis viciifolia E 35 1.04 Slovakia Oxalis stricta A 2 0.06 Slovakia Pinus nigra E 5 0.15 Slovakia Robinia pseudoacacia A 10 0.30 Slovenia Impatiens glandulifera A 11 0.87 Slovenia Impatiens parviflora A 3 0.24 Slovenia Juncus tenuis A 6 0.47 Slovenia Lupinus polyphyllus A 2 0.16 Slovenia Lycopersicon esculentum A 4 0.32 Slovenia Matricaria discoidea A 6 0.47 Spain Buddleja davidii A 3 0.07 Spain Carpobrotus edulis A 4 0.09 Spain Ceratochloa cathartica A 4 0.09 Spain Cortaderia selloana A 10 0.23 Spain Crocosmia x crocosmiiflora C 2 0.05 Spain Cuscuta campestris A 5 0.11 Spain Oenothera suaveolens A 4 0.09 Spain Onobrychis viciifolia E 26 0.60 Spain Oxalis latifolia A 4 0.09 Spain Panicum capillare A 3 0.07 Spain Parthenocissus inserta A 1 0.02 Spain Paspalum dilatatum A 59 1.35 Sweden Festuca filiformis E 1 0.26 Sweden Galium album E 5 1.28 subsp. album Sweden Matricaria discoidea A 1 0.26 Sweden Medicago sativa C 3 0.77 Sweden Noccaea perfoliata E 3 0.77 Sweden Sanguisorba minor E 1 0.26 Switzerland Schedonorus arundinaceus E 1 0.04 Switzerland Solidago canadensis A 3 0.13 Switzerland Solidago gigantea A 20 0.84 Switzerland Symphyotrichum lanceolatum A 2 0.08 Switzerland Trifolium hybridum C 1 0.04 Switzerland Veronica filiformis E 16 0.67 Ukraine Fraxinus pennsylvanica A 1 0.03 Ukraine Galinsoga quadriradiata A 1 0.03 Ukraine Grindelia squarrosa A 42 1.28 Ukraine Helianthus annuus A 1 0.03 Ukraine Heracleum mantegazzianum E 5 0.15 Ukraine Hyssopus officinalis E 1 0.03 Ukraine Saponaria officinalis E 4 0.12 Ukraine Sisymbrium altissimum E 12 0.37 Ukraine Sisyrinchium septentrionale A 13 0.40 Ukraine Solidago canadensis A 3 0.09 Ukraine Syringa vulgaris E 1 0.03 Ukraine Tanacetum macrophyllum E 2 0.06 United Kingdom Lagurus ovatus E 1 0.05 United Kingdom Lathyrus latifolius E 4 0.20 United Kingdom Lolium multiflorum E 1 0.05 United Kingdom Matricaria discoidea A 10 0.50 United Kingdom Medicago sativa C 1 0.05 United Kingdom Melilotus officinalis E 4 0.20 Appendix S10. Region Taxon Origin No. of plots % of plots Austria Lolium multiflorum E 21 0.44 Baleares * NA Bosnia and Herzegovina Medicago sativa C 2 0.82 Crete * Ageratina adenophora A 1 1.02 Croatia Stachys byzantina E 1 0.19 Czech Republic Impatiens parviflora A 70 1.34 Czech Republic Silene flos-jovis E 1 0.02 Denmark Impatiens parviflora A 3 0.16 Denmark Salix viminalis E 3 0.16 Finland Noccaea brachypetala E 3 0.70 France Carduus tenuiflorus E 17 0.09 France Lathyrus sativus C 2 0.01 France Robinia pseudoacacia A 30 0.16 Germany Atriplex hortensis E 1 0.01 Germany Impatiens glandulifera A 72 0.52 Germany Rosa rugosa A 1 0.01 Greece Cyperus sharonensis A 2 0.42 Italy Carpobrotus edulis A 2 0.07 Italy Solidago gigantea A 5 0.19 Latvia Symphoricarpos albus A 1 0.11 Lithuania Senecio leucanthemifolius E 3 0.64 Netherlands Heracleum mantegazzianum E 1 0.06 Poland Acorus calamus A 14 0.13 Poland Galinsoga quadriradiata A 2 0.02 Poland Onobrychis viciifolia E 53 0.51 Poland Symphyotrichum salignum A 2 0.02 Russia (East) Echinocystis lobata A 1 0.04 Russia (South) Bidens frondosus A 2 0.15 Sicily and Malta Xanthium orientale A 6 2.68 Slovakia Rumex patientia E 1 0.03 Slovenia Medicago sativa C 41 3.24 Spain Cyperus eragrostis A 66 1.51 Spain Paspalum distichum A 53 1.22 Switzerland Bidens frondosus A 2 0.08 Switzerland Veronica peregrina A 1 0.04 Ukraine Impatiens parviflora A 1 0.03 Ukraine Tribulus terrestris E 14 0.43 United Kingdom Mimulus guttatus A 4 0.20 Subsp. Albania * Amorpha fruticosa A 1 3.45 Albania * Onobrychis viciifolia E 1 3.45 Albania * Symphyotrichum squamatum A 2 6.90 Austria Acer negundo A 10 0.21 Austria Aesculus hippocastanum E 1 0.02 Austria Ailanthus altissima A 5 0.10 Austria Amaranthus retroflexus A 5 0.10 Austria Anthriscus cerefolium C 2 0.04 Austria Armoracia rusticana E 3 0.06 Austria Bidens frondosus A 2 0.04 Austria Brassica napus C 1 0.02 Austria Bunias orientalis E 1 0.02 Austria Calendula officinalis C 1 0.02 Austria Campanula rhomboidalis E 1 0.02 Austria Castanea sativa E 5 0.10 Austria Conringia orientalis E 1 0.02 F o r R e v i e w O n l y Austria Lycium barbarum A 1 0.02 Austria Lycopersicon esculentum A 5 0.10 Austria Mahonia aquifolium A 1 0.02 Austria Malva moschata E 6 0.13 Austria Matricaria discoidea A 1 0.02 Austria Medicago sativa C 142 2.98 subsp. sativa, nothosubsp. varia Austria Melilotus indicus E 1 0.02 Austria Narcissus poeticus E 41 0.86 subsp. poeticus, subsp. radiiflorus Austria Nepeta cataria E 1 0.02 Austria Oenothera biennis A 1 0.02 Austria Onobrychis viciifolia E 243 5.10 Austria Oxalis stricta A 16 0.34 Austria Physalis alkekengi C 2 0.04 Austria Populus x canadensis C 1 0.02 Austria Prunus cerasifera E 1 0.02 Austria Prunus cerasus A 2 0.04 Austria Prunus domestica C 13 0.27 F o r R e v i e w O n l y Belarus * Bidens frondosus A 5 11.90 Belarus * Calystegia silvatica E 3 7.14 Belarus * Echinochloa crus-galli C 1 2.38 Belarus * Epilobium ciliatum A 3 7.14 subsp. adenocaulon Belarus * Erigeron canadensis A 8 19.05 Belarus * Juncus tenuis A 1 2.38 Belarus * Mimulus guttatus A 2 4.76 Belarus * Oenothera biennis A 1 2.38 Belarus * Rumex stenophyllus E 1 2.38 Belarus * Trifolium hybridum C 1 2.38 Belgium Berteroa incana E 1 0.73 Belgium Callistephus chinensis A 2 1.46 Belgium Castanea sativa E 5 3.65 Belgium Epilobium ciliatum A 9 6.57 Belgium Erigeron canadensis A 6 4.38 Belgium Juglans regia E 1 0.73 Belgium Juncus tenuis A 5 3.65 F o r R e v i e w O n l y Bosnia and Herzegovina Papaver orientale A 1 0.41 Bosnia and Herzegovina Veronica persica A 1 0.41 Bosnia and Herzegovina Xanthium orientale A 2 0.82 Bulgaria Ailanthus altissima A 1 0.03 Bulgaria Amaranthus retroflexus A 2 0.06 Bulgaria Amorpha fruticosa A 3 0.09 Bulgaria Brassica napus C 1 0.03 Bulgaria Cuscuta campestris A 1 0.03 Bulgaria Elaeagnus angustifolia A 1 0.03 Bulgaria Erigeron annuus A 43 1.24 Bulgaria Erigeron canadensis A 18 0.52 Bulgaria Gleditsia triacanthos A 1 0.03 Bulgaria Juncus tenuis A 4 0.12 Bulgaria Lavandula angustifolia E 1 0.03 Bulgaria Medicago sativa C 7 0.20 subsp. sativa Bulgaria Quercus rubra A 2 0.06 Bulgaria Reynoutria x bohemica A 1 0.03 F o r R e v i e w O n l y Crete * Avena sativa C 1 1.02 Crete * Digitaria ciliaris A 1 1.02 Crete * Eleusine indica A 1 1.02 Crete * Lycopersicon esculentum A 1 1.02 Crete * Oxalis pes-caprae A 2 2.04 Crete * Panicum dichotomiflorum A 1 1.02 Crete * Paspalum distichum A 1 1.02 Crete * Ricinus communis A 1 1.02 Crete * Symphyotrichum squamatum A 1 1.02 Crete * Xanthium orientale A 1 1.02 Croatia Amaranthus retroflexus A 2 0.37 Croatia Ambrosia artemisiifolia A 15 2.79 Croatia Amorpha fruticosa A 3 0.56 Croatia Asclepias syriaca A 1 0.19 Croatia Bidens frondosus A 1 0.19 Croatia Carex vulpinoidea A 1 0.19 Croatia Echinocystis lobata A 9 1.68 F o r R e v i e w O n l y Croatia Veronica persica A 2 0.37 Croatia Xanthium spinosum A 7 1.30 Czech Republic Acer negundo A 2 0.04 Czech Republic Acorus calamus A 1 0.02 Czech Republic Aesculus hippocastanum E 1 0.02 Czech Republic Amaranthus powellii A 1 0.02 Czech Republic Amaranthus retroflexus A 4 0.08 Czech Republic Amorpha fruticosa A 1 0.02 Czech Republic Anaphalis margaritacea A 2 0.04 Czech Republic Asparagus officinalis E 32 0.61 Czech Republic Bidens connatus A 1 0.02 Czech Republic Bidens frondosus A 35 0.67 Czech Republic Brachypodium rupestre E 1 0.02 Czech Republic Bunias orientalis E 1 0.02 Czech Republic Campanula rapunculus E 1 0.02 Czech Republic Centaurea nigra E 1 0.02 Czech Republic Cytisus scoparius E 9 0.17 F o r R e v i e w O n l y Czech Republic Iris x germanica C 2 0.04 Czech Republic Juncus tenuis A 10 0.19 Czech Republic Lamium galeobdolon C 2 0.04 subsp. argentatum Czech Republic Lathyrus hirsutus E 1 0.02 Czech Republic Lolium multiflorum E 12 0.23 Czech Republic Lolium x hybridum C 1 0.02 Czech Republic Lupinus polyphyllus A 10 0.19 Czech Republic Lycium chinense A 1 0.02 Czech Republic Lycopersicon esculentum A 4 0.08 Czech Republic Mahonia aquifolium A 1 0.02 Czech Republic Matricaria discoidea A 41 0.79 Czech Republic Medicago sativa C 47 0.90 subsp. sativa, nothosubsp. varia Czech Republic Mimulus guttatus A 6 0.12 Czech Republic Oenothera biennis A 10 0.19 Czech Republic Onobrychis viciifolia E 53 1.02 Czech Republic Opuntia A 1 0.02 Czech Republic Oxalis dillenii A 1 0.02 F o r R e v i e w O n l y Czech Republic Sisymbrium altissimum E 3 0.06 Czech Republic Sisymbrium loeselii E 3 0.06 Czech Republic Sisymbrium strictissimum E 1 0.02 Czech Republic Solanum tuberosum A 1 0.02 Czech Republic Solidago canadensis A 23 0.44 Czech Republic Solidago gigantea A 7 0.13 Czech Republic Symphyotrichum lanceolatum A 7 0.13 Czech Republic Symphyotrichum novi-belgii A 5 0.10 Czech Republic Syringa vulgaris E 3 0.06 Czech Republic Trifolium hybridum C 123 2.36 subsp. hybridum Czech Republic Veronica filiformis E 2 0.04 Czech Republic Veronica persica A 14 0.27 Czech Republic Viola suavis E 1 0.02 Czech Republic Xanthium orientale A 1 0.02 Denmark Abies alba E 2 0.11 Denmark Acorus calamus A 2 0.11 Denmark Amelanchier spicata A 1 0.05 F o r R e v i e w O n l y Denmark Lathyrus sativus C 1 0.05 Denmark Linum austriacum E 18 0.95 Denmark Lolium multiflorum E 1 0.05 Denmark Lupinus polyphyllus A 1 0.05 Denmark Lycium barbarum A 1 0.05 Denmark Lysimachia punctata E 2 0.11 Denmark Malus toringo A 2 0.11 Denmark Malva moschata E 2 0.11 Denmark Matricaria discoidea A 3 0.16 Denmark Medicago sativa C 7 0.37 subsp. sativa, nothosubsp. varia Denmark Mimulus guttatus A 2 0.11 Denmark Onobrychis viciifolia E 3 0.16 Denmark Petasites hybridus E 1 0.05 Denmark Picea abies E 19 1.01 Denmark Picea glauca A 7 0.37 Denmark Picea sitchensis A 7 0.37 Denmark Pilosella aurantiaca E 1 0.05 F o r R e v i e w O n l y Denmark Salix x fragilis E 2 0.11 Denmark Salix x smithiana E 1 0.05 Denmark Sanguisorba minor E 15 0.80 Denmark Saponaria officinalis E 1 0.05 Denmark Senecio leucanthemifolius E 4 0.21 Denmark Solidago gigantea A 3 0.16 Denmark Sorbus aria E 1 0.05 Denmark Symphoricarpos albus A 2 0.11 Denmark Symphytum x uplandicum C 4 0.21 Denmark Syringa vulgaris E 2 0.11 Denmark Trifolium hybridum C 4 0.21 Denmark Veronica longifolia E 1 0.05 Denmark Vicia sativa E 12 0.64 Denmark Vicia villosa E 1 0.05 Denmark Zea mays A 1 0.05 Estonia Aquilegia vulgaris E 2 0.21 Estonia Lolium multiflorum E 2 0.21 F o r R e v i e w O n l y Finland Noccaea caerulescens E 4 0.93 Finland Odontites vernus E 2 0.47 Finland Potentilla intermedia E 2 0.47 Finland Potentilla thuringiaca E 1 0.23 Finland Sambucus racemosa E 1 0.23 Finland Trifolium hybridum C 74 17.21 Finland Veronica agrestis E 1 0.23 France Acacia dealbata A 3 0.02 France Acer negundo A 5 0.03 France Allium ascalonicum A 3 0.02 France Amaranthus albus A 1 0.01 France Amaranthus blitoides A 1 0.01 France Amaranthus deflexus A 1 0.01 France Amaranthus emarginatus A 1 0.01 France Amaranthus hybridus A 3 0.02 France Ambrosia artemisiifolia A 9 0.05 France Angelica archangelica E 4 0.02 F o r R e v i e w O n l y France Cedrus atlantica A 4 0.02 France Collomia grandiflora A 7 0.04 France Cotula coronopifolia A 9 0.05 France Crassula lycopodioides A 145 0.76 France Crataegus germanica E 2 0.01 France Crepis sancta E 36 0.19 France Cuscuta campestris A 1 0.01 France Cuscuta suaveolens A 6 0.03 France Cymbalaria muralis E 2 0.01 France Cyperus eragrostis A 7 0.04 France Dianthus caryophyllus E 56 0.29 France Dysphania ambrosioides A 8 0.04 France Elaeagnus angustifolia A 1 0.01 France Elodea canadensis A 1 0.01 France Epilobium ciliatum A 1 0.01 France Erigeron annuus A 53 0.28 subsp. annuus, subsp. septentrionalis France Erigeron canadensis A 108 0.57 F o r R e v i e w O n l y France Lepidium didymum A 1 0.01 France Lepidium virginicum A 12 0.06 France Lindernia dubia A 1 0.01 France Lonicera japonica A 1 0.01 France Ludwigia grandiflora A 2 0.01 France Lupinus luteus E 1 0.01 France Lycopersicon esculentum A 2 0.01 France Lysimachia punctata E 1 0.01 France Matricaria discoidea A 28 0.15 France Medicago sativa C 15 0.08 subsp. sativa, nothosubsp. varia France Oenothera biennis A 27 0.14 France Oenothera longiflora A 1 0.01 France Oenothera parviflora A 2 0.01 France Oenothera suaveolens A 2 0.01 France Oxalis dillenii A 41 0.22 France Oxalis pes-caprae A 1 0.01 France Oxalis stricta A 11 0.06 F o r R e v i e w O n l y France Rorippa austriaca E 1 0.01 France Rubus spectabilis A 16 0.08 France Rudbeckia laciniata A 4 0.02 France Rumex patientia E 2 0.01 France Rumex thyrsiflorus E 76 0.40 France Ruta graveolens E 3 0.02 France Sisyrinchium angustifolium A 1 0.01 France Sisyrinchium montanum A 1 0.01 France Solanum gracile A 1 0.01 France Solidago canadensis A 18 0.09 France Solidago gigantea A 70 0.37 France Sorghum halepense A 1 0.01 France Spartina x townsendii C 2 0.01 France Sporobolus indicus A 1 0.01 France Symphyotrichum lanceolatum A 7 0.04 France Symphyotrichum novi-belgii A 1 0.01 France Symphyotrichum parviflorum A 1 0.01 F o r R e v i e w O n l y Germany Bassia scoparia E 1 0.01 Germany Berteroa incana E 86 0.62 Germany Bidens frondosus A 14 0.10 Germany Brassica napus C 3 0.02 Germany Bunias orientalis E 12 0.09 Germany Calystegia silvatica E 1 0.01 Germany Caragana arborescens A 1 0.01 Germany Centaurea nigrescens E 6 0.04 subsp. nigrescens Germany Cerastium tomentosum E 13 0.09 Germany Corispermum intermedium E 9 0.06 Germany Cotinus coggygria E 1 0.01 Germany Cotula coronopifolia A 1 0.01 Germany Crepis nicaeensis E 1 0.01 Germany Cynodon dactylon E 6 0.04 Germany Dactylis glomerata E 1 0.01 Germany Diplotaxis tenuifolia E 7 0.05 Germany Echinops sphaerocephalus E 16 0.12 F o r R e v i e w O n l y Germany Impatiens parviflora A 13 0.09 Germany Juncus tenuis A 51 0.37 Germany Laburnum anagyroides E 1 0.01 Germany Lactuca tatarica E 1 0.01 Germany Lepidium densiflorum A 1 0.01 Germany Lepidium draba E 4 0.03 Germany Lepidium heterophyllum E 1 0.01 Germany Linum austriacum E 17 0.12 Germany Lolium multiflorum E 136 0.98 Germany Lupinus polyphyllus A 16 0.12 Germany Lycium barbarum A 1 0.01 Germany Matricaria discoidea A 31 0.22 Germany Medicago sativa C 216 1.56 subsp. sativa, nothosubsp. varia Germany Mentha x piperita C 1 0.01 Germany Mimulus guttatus A 2 0.01 Germany Narcissus poeticus E 4 0.03 subsp. poeticus, subsp. radiiflorus Germany Oenothera sp. A 27 0.19 F o r R e v i e w O n l y Germany Rubus armeniacus E 2 0.01 Germany Rudbeckia laciniata A 1 0.01 Germany Rumex triangulivalvis A 1 0.01 Germany Salix integra A 5 0.04 Germany Salsola tragus E 7 0.05 Germany Salvia officinalis E 1 0.01 Germany Sanguisorba minor E 9 0.06 Germany Sedum spurium E 3 0.02 Germany Senecio leucanthemifolius E 142 1.02 Germany Sisymbrium altissimum E 1 0.01 Germany Sisymbrium loeselii E 8 0.06 Germany Solanum tuberosum A 1 0.01 Germany Solidago canadensis A 36 0.26 Germany Solidago gigantea A 51 0.37 Germany Symphyotrichum lanceolatum A 2 0.01 Germany Symphyotrichum novi-belgii A 10 0.07 Germany Symphyotrichum parviflorum A 1 0.01 F o r R e v i e w O n l y Greece Digitaria ciliaris A 1 0.21 Greece Linum numidicum A 1 0.21 Greece Oenothera drummondii A 6 1.25 Greece Xanthium spinosum A 4 0.83 Hungary Ambrosia artemisiifolia A 18 6.59 Hungary Asclepias syriaca A 1 0.37 Hungary Bidens frondosus A 1 0.37 Hungary Erigeron annuus A 4 1.47 Hungary Erigeron canadensis A 15 5.49 Hungary Fagopyrum esculentum C 1 0.37 Hungary Hemerocallis lilioasphodelus E 2 0.73 Hungary Melissa officinalis E 1 0.37 Hungary Oenothera biennis A 2 0.73 Hungary Onobrychis viciifolia E 1 0.37 Hungary Robinia pseudoacacia A 1 0.37 Hungary Solidago canadensis A 1 0.37 Hungary Solidago gigantea A 14 5.13 F o r R e v i e w O n l y Italy Cenchrus spinifex A 12 0.45 Italy Ceratochloa cathartica A 1 0.04 Italy Crepis sancta E 30 1.11 Italy Cuscuta campestris A 1 0.04 Italy Cycloloma atriplicifolium A 1 0.04 Italy Dactyloctenium aegyptium A 1 0.04 Italy Dipsacus laciniatus E 1 0.04 Italy Dysphania ambrosioides A 3 0.11 Italy Elaeagnus angustifolia A 2 0.07 Italy Erigeron annuus A 9 0.33 Italy Erigeron bonariensis A 9 0.33 Italy Erigeron canadensis A 27 1.00 Italy Erigeron sumatrensis A 5 0.19 Italy Galinsoga parviflora A 1 0.04 Italy Helianthus tuberosus A 3 0.11 Italy Impatiens parviflora A 2 0.07 Italy Juncus tenuis A 2 0.07 F o r R e v i e w O n l y Italy Sporobolus virginicus A 23 0.85 Italy Symphyotrichum squamatum A 9 0.33 Italy Trachelium caeruleum E 2 0.07 Italy Veronica persica A 11 0.41 Italy Xanthium orientale A 52 1.93 Italy Yucca gloriosa A 2 0.07 Kosovo Medicago sativa C 2 0.77 Latvia Armoracia rusticana E 1 0.11 Latvia Asparagus officinalis E 1 0.11 Latvia Centaurea calcitrapa E 1 0.11 Latvia Epilobium ciliatum A 1 0.11 Latvia Erigeron annuus A 7 0.77 Latvia Erigeron canadensis A 15 1.64 Latvia Fagopyrum esculentum C 1 0.11 Latvia Galinsoga quadriradiata A 1 0.11 Latvia Glycine max A 1 0.11 Latvia Heracleum pubescens E 1 0.11 F o r R e v i e w O n l y Latvia Symphyotrichum salignum A 1 0.11 Lithuania Acer negundo A 1 0.21 Lithuania Acorus calamus A 3 0.64 Lithuania Agropyron dasyanthum E 5 1.07 Lithuania Epilobium ciliatum A 1 0.21 Lithuania Erigeron annuus A 9 1.92 subsp. annuus, subsp. septentrionalis Lithuania Erigeron canadensis A 27 5.77 Lithuania Galinsoga quadriradiata A 2 0.43 Lithuania Glyceria striata A 1 0.21 Lithuania Helianthus tuberosus A 1 0.21 Lithuania Heliotropium arborescens A 1 0.21 Lithuania Heracleum pubescens E 3 0.64 Lithuania Hippophae rhamnoides E 1 0.21 Lithuania Hordeum jubatum A 2 0.43 Lithuania Impatiens glandulifera A 1 0.21 Lithuania Juncus tenuis A 2 0.43 Lithuania Lactuca serriola E 1 0.21 F o r R e v i e w O n l y Luxembourg * NA Macedonia * Pistacia terebinthus E 5 7.04 Moldavia Acer negundo A 7 3.48 Moldavia Ailanthus altissima A 10 4.98 Moldavia Ambrosia artemisiifolia A 10 4.98 Moldavia Cuscuta campestris A 1 0.50 Moldavia Elaeagnus angustifolia A 1 0.50 Moldavia Erigeron annuus A 4 1.99 Moldavia Erigeron canadensis A 1 0.50 Moldavia Impatiens parviflora A 1 0.50 Moldavia Lepidium draba E 1 0.50 Moldavia Oenothera parviflora A 1 0.50 Moldavia Onobrychis viciifolia E 3 1.49 Moldavia Prunus cerasifera E 1 0.50 Moldavia Robinia pseudoacacia A 1 0.50 Montenegro * NA Netherlands Acorus calamus A 15 0.89 F o r R e v i e w O n l y Netherlands Impatiens glandulifera A 3 0.18 Netherlands Juncus tenuis A 4 0.24 Netherlands Larix kaempferi A 1 0.06 Netherlands Lepidium didymum A 1 0.06 Netherlands Lepidium draba E 2 0.12 Netherlands Lolium multiflorum E 2 0.12 Netherlands Lycium chinense A 1 0.06 Netherlands Matricaria discoidea A 37 2.20 Netherlands Medicago sativa C 7 0.42 Netherlands Oenothera sp. A 1 0.06 Netherlands Oenothera biennis A 2 0.12 Netherlands Oxalis stricta A 1 0.06 Netherlands Picea abies E 1 0.06 Netherlands Pilosella aurantiaca E 1 0.06 Netherlands Pilosella caespitosa E 1 0.06 Netherlands Pinus nigra E 1 0.06 Netherlands Plantago arenaria E 1 0.06 F o r R e v i e w O n l y Poland Aesculus hippocastanum E 4 0.04 Poland Amaranthus retroflexus A 4 0.04 Poland Anthemis ruthenica E 1 0.01 Poland Anthoxanthum aristatum E 11 0.10 Poland Artemisia austriaca E 2 0.02 Poland Asclepias syriaca A 1 0.01 Poland Atriplex tatarica E 5 0.05 Poland Beckmannia eruciformis E 10 0.10 Poland Berberis verruculosa A 1 0.01 Poland Bidens connatus A 3 0.03 Poland Bidens frondosus A 70 0.67 Poland Brassica elongata E 1 0.01 Poland Brassica napus C 2 0.02 Poland Brassica rapa E 1 0.01 Poland Bunias orientalis E 1 0.01 Poland Celastrus orbiculatus A 1 0.01 Poland Corispermum hyssopifolium E 1 0.01 F o r R e v i e w O n l y Poland Galium parisiense E 1 0.01 Poland Geranium pyrenaicum E 1 0.01 Poland Glyceria striata A 2 0.02 Poland Helianthus tuberosus A 3 0.03 Poland Heracleum mantegazzianum E 13 0.12 Poland Heracleum pubescens E 22 0.21 Poland Chenopodium strictum E 4 0.04 Poland Impatiens capensis A 2 0.02 Poland Impatiens glandulifera A 46 0.44 Poland Impatiens parviflora A 75 0.71 Poland Inula helenium E 1 0.01 Poland Juglans cinerea A 1 0.01 Poland Juncus tenuis A 66 0.63 Poland Lathyrus sativus C 1 0.01 Poland Lepidium virginicum A 1 0.01 Poland Linaria spartea E 2 0.02 Poland Linum austriacum E 1 0.01 F o r R e v i e w O n l y Poland Ornithopus sativus E 1 0.01 Poland Oxalis fontana A 26 0.25 Poland Oxalis stricta A 18 0.17 Poland Papaver somniferum E 2 0.02 Poland Phalaris paradoxa E 1 0.01 Poland Picea pungens A 1 0.01 Poland Pinguicula grandiflora E 2 0.02 Poland Pinus banksiana A 1 0.01 Poland Populus x canadensis C 1 0.01 Poland Potentilla intermedia E 1 0.01 Poland Prunus cerasifera E 1 0.01 Poland Prunus cerasus A 11 0.10 Poland Prunus serotina A 31 0.30 Poland Pseudotsuga menziesii A 1 0.01 Poland Quercus rubra A 11 0.10 Poland Reynoutria japonica A 2 0.02 Poland Rhus typhina A 1 0.01 F o r R e v i e w O n l y Poland Syringa vulgaris E 5 0.05 Poland Tanacetum parthenium E 2 0.02 Poland Thladiantha dubia A 5 0.05 Poland Trifolium hybridum C 340 3.24 subsp. elegans, subsp. hybridum Poland Trifolium incarnatum E 1 0.01 Poland Veronica filiformis E 2 0.02 Poland Veronica persica A 12 0.11 Poland Vicia grandiflora E 13 0.12 Poland Vicia pannonica E 1 0.01 Poland x Festulolium loliaceum C 1 0.01 Poland Xanthium sp. A 2 0.02 Poland Xanthium orientale A 12 0.11 Portugal Arctotheca calendula A 2 1.25 Portugal Carpobrotus edulis A 30 18.75 Portugal Hydrocotyle bonariensis A 5 3.13 Portugal Oxalis debilis A 1 0.63 Portugal Paspalum dilatatum A 1 0.63 F o r R e v i e w O n l y Russia (East) Erigeron canadensis A 25 1.07 Russia (East) Grindelia squarrosa A 2 0.09 Russia (East) Helianthus annuus A 2 0.09 Russia (East) Heracleum pubescens E 4 0.17 Russia (East) Leontodon biscutellifolius E 1 0.04 Russia (East) Lepidium densiflorum A 1 0.04 Russia (East) Lupinus polyphyllus A 9 0.39 Russia (East) Matricaria discoidea A 7 0.30 Russia (East) Medicago sativa C 35 1.50 Russia (East) Myosotis alpestris E 1 0.04 Russia (East) Oenothera biennis A 3 0.13 Russia (East) Onobrychis viciifolia E 3 0.13 Russia (East) Parthenocissus quinquefolia A 1 0.04 Russia (East) Physocarpus opulifolius A 1 0.04 Russia (East) Potentilla grisea A 8 0.34 Russia (East) Prunus cerasus A 3 0.13 Russia (East) Sambucus racemosa E 5 0.21 F o r R e v i e w O n l y Russia (South) Bidens x garumnae C 2 0.15 Russia (South) Cuscuta campestris A 2 0.15 Russia (South) Diplotaxis tenuifolia E 1 0.08 Russia (South) Elaeagnus angustifolia A 1 0.08 Russia (South) Erigeron canadensis A 5 0.39 Russia (South) Fraxinus pennsylvanica A 4 0.31 Russia (South) Helianthus lenticularis A 3 0.23 Russia (South) Lepidium densiflorum A 1 0.08 Russia (South) Medicago sativa C 3 0.23 Russia (South) Oenothera biennis A 2 0.15 Russia (South) Onobrychis viciifolia E 1 0.08 Russia (South) Prunus armeniaca A 1 0.08 Russia (South) Sisymbrium orientale E 1 0.08 Russia (South) Xanthium orientale A 13 1.00 subsp. californicum, subsp. riparium Russia (South) Xanthium spinosum A 2 0.15 Russia (South) Xanthium strumarium E 90 6.96 Romania Alcea rosea C 1 0.02 F o r R e v i e w O n l y Slovakia Acorus calamus A 1 0.03 Slovakia Achillea macrophylla E 1 0.03 Slovakia Amaranthus retroflexus A 2 0.06 Slovakia Anthriscus cerefolium C 1 0.03 Slovakia Bidens frondosus A 6 0.18 Slovakia Bunias orientalis E 2 0.06 Slovakia Datura stramonium A 1 0.03 Slovakia Echinops sphaerocephalus E 1 0.03 Slovakia Epilobium ciliatum A 6 0.18 Slovakia Erechtites hieraciifolius A 1 0.03 Slovakia Erigeron annuus A 18 0.53 subsp. annuus, subsp. septentrionalis Slovakia Erigeron canadensis A 12 0.36 Slovakia Erucastrum gallicum E 1 0.03 Slovakia Galinsoga parviflora A 3 0.09 Slovakia Galinsoga quadriradiata A 3 0.09 Slovakia Galium verrucosum E 1 0.03 Slovakia Geranium pyrenaicum E 1 0.03 F o r R e v i e w O n l y Slovakia Rumex thyrsiflorus E 8 0.24 Slovakia Sedum rupestre E 1 0.03 Slovakia Sisymbrium austriacum E 1 0.03 Slovakia Solidago canadensis A 3 0.09 Slovakia Solidago gigantea A 3 0.09 Slovakia Symphyotrichum lanceolatum A 1 0.03 Slovakia Symphyotrichum novi-belgii A 4 0.12 Slovakia Syringa vulgaris E 1 0.03 Slovakia Trifolium hybridum C 46 1.37 Slovakia Veronica filiformis E 1 0.03 Slovakia Veronica peregrina A 2 0.06 Slovakia Veronica persica A 3 0.09 Slovakia Viola suavis E 1 0.03 Slovakia Xanthium orientale A 1 0.03 Slovenia Ailanthus altissima A 1 0.08 Slovenia Amaranthus cruentus A 2 0.16 Slovenia Amaranthus retroflexus A 1 0.08 F o r R e v i e w O n l y Slovenia Oenothera biennis A 2 0.16 Slovenia Oxalis stricta A 10 0.79 Slovenia Panicum dichotomiflorum A 1 0.08 Slovenia Reynoutria japonica A 2 0.16 Slovenia Robinia pseudoacacia A 3 0.24 Slovenia Rudbeckia laciniata A 12 0.95 Slovenia Scirpus georgianus A 2 0.16 Slovenia Sedum spurium E 1 0.08 Slovenia Solidago canadensis A 11 0.87 Slovenia Solidago gigantea A 47 3.71 Slovenia Tagetes erecta A 1 0.08 Slovenia Veronica persica A 12 0.95 Slovenia Viola cornuta E 4 0.32 Slovenia x Festulolium sp. C 1 0.08 Slovenia Xanthium orientale A 2 0.16 Spain Aesculus hippocastanum E 1 0.02 Spain Amaranthus albus A 1 0.02 F o r R e v i e w O n l y Spain Delairea odorata A 1 0.02 Spain Dysphania ambrosioides A 2 0.05 Spain Eleocharis bonariensis A 1 0.02 Spain Erigeron bonariensis A 4 0.09 Spain Erigeron canadensis A 33 0.76 Spain Erigeron floribundus A 19 0.44 Spain Galega officinalis E 1 0.02 Spain Galinsoga quadriradiata A 1 0.02 Spain Gnaphalium coarctatum A 4 0.09 Spain Helianthus x laetiflorus C 4 0.09 Spain Hydrangea macrophylla A 1 0.02 Spain Hypericum hircinum E 1 0.02 Spain Impatiens balfourii A 5 0.11 Spain Ipomoea indica A 1 0.02 Spain Iris x germanica C 2 0.05 Spain Juncus tenuis A 4 0.09 Spain Lepidium didymum A 1 0.02 F o r R e v i e w O n l y Spain Paspalum vaginatum A 8 0.18 Spain Persicaria pensylvanica A 1 0.02 Spain Pinus radiata A 1 0.02 Spain Pinus strobus A 5 0.11 Spain Populus x canadensis C 1 0.02 Spain Prunus x fruticans E 1 0.02 Spain Reynoutria japonica A 1 0.02 Spain Robinia pseudoacacia A 3 0.07 Spain Selaginella kraussiana A 1 0.02 Spain Solanum chenopodioides A 4 0.09 Spain Solanum tuberosum A 1 0.02 Spain Sorghum halepense A 2 0.05 Spain Sporobolus indicus A 13 0.30 Spain Stenotaphrum secundatum A 9 0.21 Spain Symphyotrichum squamatum A 67 1.54 Spain Tradescantia fluminensis A 2 0.05 Spain Tropaeolum majus A 1 0.02 F o r R e v i e w O n l y Switzerland Bromopsis inermis E 7 0.29 Switzerland Ceratochloa carinata A 1 0.04 Switzerland Crepis nicaeensis E 1 0.04 Switzerland Dipsacus sativus C 2 0.08 Switzerland Erigeron annuus A 35 1.47 Switzerland Erigeron canadensis A 3 0.13 Switzerland Euphorbia lathyris E 1 0.04 Switzerland Gaultheria procumbens A 1 0.04 Switzerland Helianthus tuberosus A 1 0.04 Switzerland Helminthotheca echioides E 1 0.04 Switzerland Hesperis matronalis E 1 0.04 Switzerland Impatiens glandulifera A 10 0.42 Switzerland Impatiens parviflora A 2 0.08 Switzerland Juncus tenuis A 5 0.21 Switzerland Lolium multiflorum E 5 0.21 Switzerland Lonicera japonica A 1 0.04 Switzerland Matricaria discoidea A 1 0.04 F o r R e v i e w O n l y Switzerland Veronica persica A 32 1.35 Switzerland Vicia lutea E 5 0.21 Switzerland Vicia sativa E 1 0.04 Switzerland Vicia villosa E 8 0.34 subsp. varia, subsp. villosa Ukraine Acer negundo A 8 0.24 Ukraine Acorus calamus A 5 0.15 Ukraine Ailanthus altissima A 3 0.09 Ukraine Allium ascalonicum A 4 0.12 Ukraine Amaranthus albus A 3 0.09 Ukraine Amaranthus retroflexus A 1 0.03 Ukraine Ambrosia artemisiifolia A 21 0.64 Ukraine Anchusa stylosa E 9 0.27 Ukraine Anthemis odontostephana A 1 0.03 Ukraine Anthericum liliago E 2 0.06 Ukraine Artemisia annua E 1 0.03 Ukraine Asclepias syriaca A 2 0.06 Ukraine Bidens frondosus A 3 0.09 F o r R e v i e w O n l y Ukraine Iva xanthiifolia A 2 0.06 Ukraine Juncus tenuis A 21 0.64 Ukraine Lathyrus sativus C 1 0.03 Ukraine Lepidium densiflorum A 3 0.09 Ukraine Lolium multiflorum E 1 0.03 Ukraine Lupinus polyphyllus A 3 0.09 Ukraine Lycium barbarum A 3 0.09 Ukraine Medicago sativa C 16 0.49 Ukraine Myagrum perfoliatum E 1 0.03 Ukraine Oenothera biennis A 18 0.55 Ukraine Oenothera rubricaulis A 14 0.43 Ukraine Onobrychis viciifolia E 53 1.62 Ukraine Oxalis fontana A 5 0.15 Ukraine Oxalis stricta A 2 0.06 Ukraine Papaver somniferum E 1 0.03 Ukraine Parthenocissus inserta A 1 0.03 Ukraine Petrorhagia saxifraga E 1 0.03 F o r R e v i e w O n l y Ukraine Trifolium hybridum C 73 2.23 subsp. hybridum Ukraine Ulmus pumila A 1 0.03 Ukraine Veronica peregrina A 1 0.03 Ukraine Veronica persica A 5 0.15 Ukraine Xanthium orientale A 34 1.04 subsp. italicum, subsp. riparium Ukraine Xanthium strumarium E 3 0.09 subsp. sibiricum United Kingdom Acer pseudoplatanus E 9 0.45 United Kingdom Alchemilla conjuncta E 1 0.05 United Kingdom Armoracia rusticana E 1 0.05 United Kingdom Brassica napus C 1 0.05 United Kingdom Briza maxima E 2 0.10 United Kingdom Bromus hordeaceus E 13 0.64 United Kingdom Bromus lepidus E 1 0.05 United Kingdom Centaurea aspera E 2 0.10 United Kingdom Centaurea calcitrapa E 1 0.05 United Kingdom Claytonia perfoliata A 1 0.05 United Kingdom Claytonia sibirica A 1 F o r R e v i e w O n l y United Kingdom Muscari neglectum E 1 0.05 United Kingdom Myrrhis odorata E 2 0.10 United Kingdom Oenothera stricta A 1 0.05 United Kingdom Pentaglottis sempervirens E 1 0.05 United Kingdom Pinus nigra E 1 0.05 United Kingdom Potentilla recta E 1 0.05 United Kingdom Reynoutria japonica A 1 0.05 United Kingdom Ribes nigrum E 1 0.05 United Kingdom Senecio squalidus E 2 0.10 United Kingdom Sinapis alba E 1 0.05 United Kingdom Solidago canadensis A 1 0.05 United Kingdom Symphoricarpos albus A 1 0.05 United Kingdom Trifolium hybridum C 1 0.05 United Kingdom Veronica persica A 14 0.69 United Kingdom Veronica praecox E 1 0.05 United Kingdom x Festulolium loliaceum C 6 0.30 306 F o r R e v i e w O n l y 0.05 287 Heavy-metal grassland in Western and Central Europe 93 93 0 0 0 0 0.0 0.0 9 9 0 0.0 0.0 R21 Mesic permanent pasture of lowlands and mountains 2632 2506 126 67 49 10 4.8 2.5 6542 5690 852 13.0 9.8 R22 Low and medium altitude hay meadow 3184 2977 207 82 110 15 6.5 2.6 22885 19570 3315 14.5 7.4 R52 Forest fringe of acidic nutrient-poor 491 480 11 8 3 0 2.2 1.6 305 290 15 4.9 3.9 Austria Echium italicum E 1 0.02 Austria Epilobium ciliatum A 10 0.21 Austria Robinia pseudoacacia A 10 0.21 Austria Rudbeckia laciniata A 4 0.08 Belgium Lathyrus latifolius E 1 0.73 Belgium Melilotus albus E 1 0.73 Bulgaria Robinia pseudoacacia A 1 0.03 Bulgaria Setaria italica A 2 0.06 Croatia Erigeron annuus A 145 27.00 Croatia Erigeron canadensis A 11 2.05 Czech Republic Datura stramonium A 1 0.02 Czech Republic Digitalis purpurea E 1 0.02 Czech Republic Oxalis stricta A 5 0.10 Czech Republic Peucedanum austriacum E 1 0.02 Denmark Berberis thunbergii A 1 0.05 Denmark Berberis vulgaris E 7 0.37 Denmark Pinus contorta A 1 0.05 Denmark Pinus mugo E 7 0.37 Estonia Lolium perenne E 1 0.10 Estonia Lotus maritimus E 2 0.21 France Armoracia rusticana E 1 0.01 France Artemisia biennis A 1 0.01 France Erigeron sumatrensis A 5 0.03 France Erysimum cheiri E 2 0.01 France Panicum capillare A 9 0.05 France Paspalum dilatatum A 3 0.02 France Symphyotrichum salignum A 3 0.02 France Symphyotrichum squamatum A 2 0.01 Germany Elaeagnus commutata A 1 0.01 Germany Epilobium ciliatum A 72 0.52 Germany Oenothera biennis A 31 0.22 Germany Onobrychis viciifolia E 177 1.28 Germany Symphyotrichum salignum A 2 0.01 Germany Symphytum asperum E 1 0.01 Hungary Xanthium sp. A 1 0.37 Ireland Crepis vesicaria E 2 1.36 Italy Matricaria discoidea A 3 0.11 Italy Oenothera biennis A 40 1.49 Latvia Impatiens glandulifera A 2 0.22 Latvia Impatiens parviflora A 1 0.11 Lithuania Lupinus polyphyllus A 8 1.71 Lithuania Matricaria discoidea A 22 4.70 Netherlands Amelanchier lamarckii A 3 0.18 Netherlands Aronia x prunifolia A 1 0.06 Netherlands Populus alba E 1 0.06 Netherlands Prunus serotina A 17 1.01 Poland Corispermum intermedium E 2 0.02 Poland Cornus sericea A 2 0.02 Poland Lolium multiflorum E 22 0.21 Poland Lupinus luteus E 5 0.05 Poland Robinia pseudoacacia A 11 0.10 Poland Rosa glauca E 2 0.02 Portugal Paspalum distichum A 2 1.25 Russia (Central) * Lepidium draba E 1 2.38 Russia (East) Symphyotrichum salignum A 1 0.04 Russia (East) Trifolium pratense C 2 0.09 Romania Alchemilla conjuncta E 3 0.06 Romania Amaranthus albus A 3 0.06 Slovakia Heracleum mantegazzianum E 1 0.03 Slovakia Hordeum marinum E 3 0.09 Slovenia Ambrosia artemisiifolia A 6 0.47 Slovenia Bidens frondosus A 1 0.08 Spain Amaranthus bouchonii A 1 0.02 Spain Amaranthus hybridus A 1 0.02 Spain Lycopersicon esculentum A 1 0.02 Spain Malcolmia maritima E 1 0.02 Spain Verbena bonariensis A 1 0.02 Spain Veronica persica A 9 0.21 Switzerland Oenothera biennis A 1 0.04 Switzerland Oenothera parviflora A 1 0.04 Ukraine Cannabis sativa E 1 0.03 Ukraine Carthamus lanatus E 2 0.06 Ukraine Plagiorhegma dubium A 2 0.06 Ukraine Potentilla indica A 1 0.03 United Kingdom Consolida ajacis E 1 0.05 United Kingdom Cotoneaster microphyllus A 2 288 0.10 soils R55 Austria Erigeron annuus A 242 Austria Rumex cristatus E 3 0.06 Belgium Oenothera parviflora A 1 0.73 Bulgaria Solidago canadensis A 1 0.03 Croatia Galinsoga parviflora A 2 0.37 Czech Republic Dipsacus strigosus E 1 0.02 Czech Republic Peucedanum ostruthium E 10 0.19 Denmark Brassica napus C 2 0.11 Denmark Pinus nigra E 2 0.11 Estonia Medicago sativa C 3 0.31 France Artemisia verlotiorum A 1 0.01 France Euthamia graminifolia A 1 0.01 France Paspalum distichum A 6 0.03 France Tanacetum coccineum E 1 0.01 Germany Eragrostis minor E 1 0.01 Germany Ornithogalum nutans E 1 0.01 Germany Syringa vulgaris E 2 0.01 Ireland Epilobium brunnescens A 5 3.40 Italy Oenothera glazioviana A 2 0.07 Latvia Juncus tenuis A 1 0.11 Lithuania Medicago sativa C 1 0.21 Netherlands Azolla filiculoides A 1 0.06 Netherlands Quercus rubra A 3 0.18 Poland Cymbalaria muralis E 1 0.01 Poland Lupinus polyphyllus A 79 0.75 Poland Rosa rugosa A 3 0.03 Russia (Central) * Marrubium peregrinum E 8 19.05 Russia (East) Ulmus pumila A 1 0.04 Romania Amaranthus hybridus A 1 0.02 Slovakia Chenopodium strictum E 3 0.09 Slovenia Echinocystis lobata A 18 1.42 Spain Amaranthus retroflexus A 1 0.02 Spain Melissa officinalis E 5 0.11 Spain Xanthium orientale A 4 0.09 subsp. orientale, subsp. italicum Switzerland Onobrychis viciifolia E 109 4.59 Ukraine Cirsium ciliatum E 1 0.03 Ukraine Prunus armeniaca A 3 0.09 United Kingdom Crepis vesicaria E 4 0.20 5.08 subsp. annuus, subsp. septentrionalis Lowland moist or wet tall-herb and fern 1913 1735 178 117 49 12 9.3 6.1 4723 3638 1085 23.0 20.5 Austria Erigeron canadensis A 27 0.57 Austria Sedum spurium E 1 Belgium Oxalis stricta A 1 0.73 Bulgaria Trigonella caerulea A 2 0.06 Croatia Helianthus tuberosus A 1 0.19 Czech Republic Echinocystis lobata A 4 0.08 Czech Republic Picea pungens A 4 0.08 Denmark Bromopsis inermis E 1 0.05 Denmark Populus alba E 1 0.05 Estonia Trifolium alpestre E 2 0.21 France Arundo donax A 4 0.02 France Fragaria ananassa C 1 0.01 France Paspalum vaginatum A 2 0.01 France Veronica filiformis E 2 0.01 Germany Erigeron annuus A 19 0.14 subsp. annuus, subsp. septentrionalis Germany Ornithopus sativus E 5 0.04 Germany Telekia speciosa E 19 0.14 Ireland Lepidium didymum A 2 1.36 Italy Oenothera parviflora A 1 0.04 Latvia Lupinus polyphyllus A 4 0.44 Lithuania Mimulus guttatus A 1 0.21 Netherlands Berteroa incana E 1 0.06 Netherlands Rumex thyrsiflorus E 23 1.37 Poland Diplotaxis muralis E 1 0.01 Poland Lycium barbatum A 1 0.01 Poland Rubia tinctorium E 2 0.02 Russia (Caucasus) Ailanthus altissima A 1 0.35 Russia (East) Verbascum phlomoides E 3 0.13 Romania Amaranthus retroflexus A 7 0.14 Slovakia Impatiens glandulifera A 3 0.09 Slovenia Erechtites hieraciifolius A 1 0.08 Spain Artemisia verlotiorum A 2 0.05 Spain Mentha spicata C 7 0.16 Spain Xanthium spinosum A 2 0.05 Switzerland Opuntia humifusa A 1 0.04 Ukraine Cuscuta campestris A 7 0.21 Ukraine Prunus cerasifera E 1 0.03 United Kingdom Cymbalaria muralis E 1 0.05 0.02 fringe Austria Fumaria capreolata E 1 0.02 Austria Senecio leucanthemifolius E 1 0.02 Belgium Parthenocissus inserta A 1 0.73 Bulgaria Xanthium orientale A 5 0.14 subsp. italicum, subsp. riparium Croatia Impatiens glandulifera A 6 1.12 Czech Republic Echinops sphaerocephalus E 19 0.36 Czech Republic Pinus nigra E 6 0.12 Denmark Centaurea nigra E 5 0.27 Denmark Populus x canescens E 1 0.05 Finland Achillea ptarmica E 170 39.53 France Asclepias syriaca A 1 0.01 France Freesia refracta A 2 0.01 France Phaseolus vulgaris A 1 0.01 France Veronica peregrina A 3 0.02 Germany Erigeron canadensis A 325 2.34 Germany Oxalis stricta A 1 0.01 Germany Thymus pulegioides E 40 0.29 Ireland Matricaria discoidea A 3 2.04 Italy Oenothera suaveolens A 2 0.07 Latvia Malva sylvestris E 2 0.22 Lithuania Oenothera ammophila A 1 0.21 Netherlands Bidens frondosus A 3 0.18 Netherlands Senecio inaequidens A 5 0.30 Poland Echinocystis lobata A 12 0.11 Poland Lycopersicon esculentum A 1 0.01 Poland Rudbeckia laciniata A 8 0.08 Russia (Caucasus) Ambrosia artemisiifolia A 3 1.04 Russia (East) Xanthium orientale A 6 0.26 Romania Ambrosia artemisiifolia A 7 0.14 Slovakia Impatiens parviflora A 8 0.24 Slovenia Erigeron annuus A 167 13.18 subsp. annuus, subsp. septentrionalis Spain Arundo donax A 3 0.07 Spain Mentha x piperita C 1 0.02 Spain Yucca gloriosa A 1 0.02 Switzerland Ornithogalum nutans E 1 0.04 Ukraine Cuscuta epilinum C 1 0.03 Ukraine Prunus domestica C 2 0.06 United Kingdom Epilobium ciliatum A 3 0.15 0.7 R1D Mediterranean closely grazed dry grassland 864 860 4 2 2 0 0.5 0.2 149 125 24 16.1 15.4 R23 Mountain hay meadow 1387 1374 13 5 6 2 0.9 0.4 1400 1366 34 2.4 0.6 R24* Iberian summer pasture (vallicar) 114 114 0 0 0 0 0.0 0.0 15 15 0 0.0 R56 Montane to subalpine moist or wet tall-herb and fern fringe 1148 1146 2 1 1 0 0.2 0.1 832 830 2 0.2 Austria Galinsoga quadriradiata A 2 0.04 Austria Sinapis alba E 1 0.02 Belgium Picea abies E 1 0.73 Bulgaria Xanthium spinosum A 2 0.06 Croatia Matricaria discoidea A 5 0.93 Czech Republic Epilobium ciliatum A 91 1.74 Czech Republic Populus x canadensis C 1 0.02 Denmark Cerastium tomentosum E 1 0.05 Denmark Potentilla recta E 2 0.11 Finland Cota triumfettii E 2 0.47 France Atriplex tatarica E 1 0.01 France Galega officinalis E 13 0.07 France Populus x canadensis C 2 0.01 France Veronica persica A 32 0.17 Germany Fritillaria meleagris E 25 0.18 Germany Pinus nigra E 4 0.03 Germany Trifolium hybridum C 131 0.94 subsp. hybridum Ireland Rhododendron ponticum E 1 0.68 Italy Onobrychis viciifolia E 129 4.79 Latvia Medicago sativa C 29 3.18 Lithuania Oenothera biennis A 14 2.99 Netherlands Brassica napus C 1 0.06 Netherlands Solanum tuberosum A 1 0.06 Poland Echinops sphaerocephalus E 5 0.05 Poland Malva moschata E 4 0.04 Poland Rumex confertus E 20 0.19 Russia (Caucasus) Amorpha fruticosa A 2 0.69 Russia (Kaliningrad) * Epilobium ciliatum A 1 33.33 Romania Amorpha fruticosa A 5 0.10 Slovakia Juncus tenuis A 17 0.51 Slovenia Erigeron canadensis A 27 2.13 Spain Avena sativa C 3 0.07 Spain Mirabilis jalapa A 1 0.02 Spain Zantedeschia aethiopica A 3 0.07 Switzerland Panicum capillare A 4 0.17 Ukraine Dasypyrum villosum E 2 0.06 Ukraine Quercus rubra A 2 0.06 United Kingdom Erica scoparia E 1 0.05 0.1 Austria Galium verrucosum E 1 Austria Sisymbrium altissimum E 1 0.02 Belgium Populus alba E 1 0.73 Corsica Carpobrotus edulis A 1 0.37 Croatia Medicago sativa C 25 4.66 Czech Republic Erigeron annuus A 20 0.38 Czech Republic Prunus armeniaca A 1 0.02 Denmark Clematis vitalba E 1 0.05 Denmark Prunus cerasifera E 60 3.18 Finland Epilobium ciliatum A 30 6.98 France Avena sativa C 1 0.01 France Galinsoga parviflora A 2 0.01 France Potentilla indica A 1 0.01 France Xanthium orientale A 2 0.01 Germany Galinsoga parviflora A 10 0.07 Germany Populus balsamifera A 1 0.01 Germany Trifolium resupinatum E 1 0.01 Ireland Veronica persica A 2 1.36 Italy Oxalis dillenii A 1 0.04 Latvia Oenothera biennis A 14 1.54 Lithuania Oenothera rubricaulis A 7 1.50 Netherlands Claytonia perfoliata A 1 0.06 Netherlands Solidago gigantea A 5 0.30 Poland Epilobium ciliatum A 26 0.25 subsp. adenocaulon, subsp. ciliatum Poland Matricaria discoidea A 68 0.65 Poland Salix daphnoides E 1 0.01 Russia (Caucasus) Diplotaxis tenuifolia E 1 0.35 Russia (Kaliningrad) * Lupinus polyphyllus A 1 33.33 Romania Aristolochia macrophylla A 1 0.02 Slovakia Lolium multiflorum E 3 0.09 Slovenia Galinsoga parviflora A 9 0.71 Spain Baccharis halimifolia A 1 0.02 Spain Oenothera biennis A 9 0.21 Sweden Berberis vulgaris E 1 0.26 Switzerland Potentilla indica A 1 0.04 Ukraine Echinocystis lobata A 1 0.03 Ukraine Raphanus sativus C 1 0.03 United Kingdom Fragaria ananassa C 1 0.05 0.02 0.0 R61* Mediterranean inland salt steppe 146 144 2 2 0 0 1.4 1.4 79 75 4 5.1 Austria Geranium pyrenaicum E 27 0.57 Austria Solidago canadensis A 15 0.31 Belgium Populus x canadensis C 1 0.73 Corsica Cotula coronopifolia A 6 2.24 Croatia Oenothera biennis A 1 0.19 Czech Republic Erigeron canadensis A 40 0.77 Czech Republic Reynoutria A 3 0.06 Denmark Cotoneaster horizontalis A 3 0.16 Denmark Prunus serotina A 14 0.74 Finland Fragaria moschata E 3 0.70 France Avena strigosa E 1 0.01 France Galinsoga quadriradiata A 3 0.02 France Prunus cerasifera E 1 0.01 France Zinnia elegans A 1 0.01 Germany Galinsoga quadriradiata A 5 0.04 Germany Populus trichocarpa A 1 0.01 Germany Ulex europaeus E 1 0.01 Italy Agave americana A 1 0.04 Italy Oxalis pes-caprae A 1 0.04 Latvia Oenothera rubricaulis A 1 0.11 Lithuania Onobrychis viciifolia E 4 0.85 Netherlands Cyanus segetum C 1 0.06 Netherlands Veronica filiformis E 1 0.06 Poland Eragrostis minor E 1 0.01 Poland Medicago sativa C 66 0.63 subsp. sativa, nothosubsp. varia Poland Senecio leucanthemifolius E 55 0.52 Russia (Caucasus) Erigeron canadensis A 1 0.35 Russia (Kaliningrad) * Rosa rugosa A 1 33.33 Romania Bidens frondosus A 5 0.10 Slovakia Malva moschata E 3 0.09 Slovenia Galinsoga quadriradiata A 4 0.32 Spain Bidens aureus A 6 0.14 Spain Oenothera fallax A 5 0.11 Sweden Bromopsis erecta E 2 0.51 Switzerland Robinia pseudoacacia A 4 0.17 Ukraine Elaeagnus angustifolia A 18 0.55 Ukraine Rapistrum rugosum E 3 0.09 United Kingdom Hesperis matronalis E 2 0.10 5.1 R62 Continental inland salt steppe 500 490 10 8 1 1 2.0 1.6 460 425 35 7.6 Austria Glebionis segetum E 1 Austria Solidago gigantea A 145 3.04 Belgium Prunus serotina A 5 3.65 Corsica Crassula lycopodioides A 4 1.49 Croatia Oxalis dillenii A 1 0.19 Czech Republic Galinsoga parviflora A 1 0.02 Czech Republic Reynoutria japonica A 7 0.13 Denmark Dactylis glomerata E 2 0.11 Denmark Reseda lutea E 4 0.21 Finland Galium album E 100 23.26 subsp. album France Baccharis halimifolia A 6 0.03 France Helianthus tuberosus A 4 0.02 France Prunus cerasus A 4 0.02 Germany Acorus calamus A 22 0.16 Germany Geranium purpureum E 1 0.01 Germany Populus x canadensis C 6 0.04 Germany Veronica filiformis E 3 0.02 Italy Ailanthus altissima A 2 0.07 Italy Oxalis stricta A 1 0.04 Latvia Pilosella aurantiaca E 1 0.11 Lithuania Parthenocissus quinquefolia A 1 0.21 Netherlands Cynodon dactylon E 4 0.24 Netherlands Veronica persica A 3 0.18 Poland Erigeron annuus A 109 1.04 subsp. annuus, subsp. septentrionalis Poland Mentha spicata C 1 0.01 Poland Silene flos-jovis E 5 0.05 Russia (Caucasus) Populus carolinensis A 1 0.35 Russia (North) * NA Romania Bidens vulgatus A 1 0.02 Slovakia Matricaria discoidea A 47 1.40 Slovenia Geum aleppicum E 4 0.32 Spain Bidens frondosus A 6 0.14 Spain Oenothera glazioviana A 1 0.02 Sweden Cota tinctoria E 3 0.77 Switzerland Rubus armeniacus E 3 0.13 Ukraine Erigeron annuus A 150 4.58 Ukraine Rhus typhina A 1 0.03 United Kingdom Impatiens capensis A 1 0.05 0.02 7.2 Austria Hemerocallis lilioasphodelus E 6 0.13 Austria Symphyotrichum lanceolatum A 7 0.15 Belgium Quercus rubra A 2 1.46 Corsica Cyperus eragrostis A 1 0.37 Croatia Oxalis stricta A 2 0.37 Czech Republic Galinsoga quadriradiata A 8 0.15 Czech Republic Reynoutria sachalinensis A 1 0.02 Denmark Digitalis purpurea E 1 0.05 Denmark Reynoutria japonica A 1 0.05 Finland Galium x pomeranicum E 5 1.16 France Berteroa incana E 98 0.51 France Impatiens balfourii A 2 0.01 France Prunus domestica C 1 0.01 Germany Aesculus hippocastanum E 1 0.01 Germany Geranium pyrenaicum E 10 0.07 Germany Prunus cerasus A 7 0.05 Germany Veronica persica A 16 0.12 Italy Ambrosia psilostachya A 18 0.67 Italy Paspalum distichum A 20 0.74 Latvia Plantago arenaria E 1 0.11 Lithuania Petasites hybridus E 7 1.50 Netherlands Echinochloa crus-galli C 1 0.06 Netherlands x Festulolium loliaceum C 1 0.06 Poland Erigeron canadensis A 291 2.77 Poland Mimulus guttatus A 6 0.06 Poland Sisymbrium loeselii E 8 0.08 Russia (Caucasus) Spartium junceum E 1 0.35 Russia (NorthWest) Rosa rugosa A 1 2.27 Romania Cuscuta campestris A 8 0.16 Slovakia Medicago sativa C 35 1.04 subsp. sativa, nothosubsp. varia Slovenia Helianthus tuberosus A 4 0.32 Spain Bromopsis inermis E 1 0.02 Spain Oenothera rosea A 7 0.16 Sweden Cynoglossum officinale E 2 0.51 Switzerland Salix repens E 2 0.08 Ukraine Erigeron canadensis A 41 1.25 Ukraine Robinia pseudoacacia A 3 0.09 United Kingdom Impatiens glandulifera A 9 0.45 307

Journal of Vegetation Science

Acknowledgements

Akademie VÄ›d ÄOEeské Republiky, (Grant / Award Number: 'RVO 67985939 ') Comunidad de Madrid, (Grant / Award Number: 'S2018/EMT-4338') Eusko Jaurlaritza, (Grant / Award Number: 'IT936-16') Grantová Agentura ÄOEeské Republiky, (Grant / Award Number: '19-28491X','19-28807X ') Russian Academy of Sciences, (Grant / Award Number: 'ÐÐÐÐ-Ð19-119031290052-1') Villum Fonden, (Grant / Award Number: '16549') Funding information 62 IA, VK, JD, MV, IK and MC were funded by the Czech Science Foundation, programme EXPRO (project no. 19-28491X). 63 PP and JP were supported by EXPRO grant no. 19-28807X (Czech Science Foundation) and long-term research 64 development project RVO 67985939 (Czech Academy of Sciences). IB was supported by the Basque Government 65 (IT936-16). AL was supported by Komarov Botanical Institute, RAS (АААА-А19-119031290052-1). JCS considers this 66 work a contribution to his VILLUM Investigator project "Biodiversity Dynamics in a Changing World" funded by 67 VILLUM FONDEN (grant 16549). RGG was supported by REMEDINAL network (S2018/EMT-4338).

FloraCatalana.net. La flora del nostre entorn. Available at http://www.floracatalana.net; [accessed June 2020] Flora Nordica online. Available at http://www.floranordica.org/ [accessed June 2020] Flora of Greece web. Vascular plants of Greece: An annotated checklist. Available at http://portal.cybertaxonomy.org/flora-greece [accessed June 2020] Flora of North America. Available at http://beta.floranorthamerica.org [accessed June 2020] Flora Vascular de Andalucía Occidental. Available at www.floravascular.com[accessed June 2020] French Flora database (baseflor), project of Flore et végétation de la France et du Monde: CATMINAT. Available at http://philippe.julve.pagespersoorange.fr/catminat.htm [accessed June 2020] GreekFlora.gr. Available at https://www.greekflora.gr/ [accessed June 2020] Guarino, R., La Rosa, M. and Pignatti, S. (Eds) (2019) Flora d'Italia, volume 4. Horváth, F., Dobolyi, Z. K., Morschhauser, T., Lõkös, L., Karas, L. and Szerdahelyi, T. (1995) FLÓRA adatbázis 1.2 -taxonlista és attribútum-állomány. FLÓRA munkacsoport, MTA-ÖBKI, MTM Növénytára, Vácrátót, p: 252. Klotz, S., Kühn, I. and Durka, W. (2002) BIOLFLOR -Eine Datenbank zu biologischökologischen Merkmalen der Gefäßpflanzen in Deutschland. Schriftenreihe für Vegetationskunde, 38, 1-334. Natura Bohemica. Available at http://www.naturabohemica.cz [accessed June 2020] NOBANIS. European Network on Invasive Alien Species. Gateway to Information on Invasive Alien species in North and Central Europe. Available at www.nobanis.org [accessed June 2020] Pladias. Database of the Czech flora and vegetation. Available at www.pladias.cz [accessed June 2020]

data. IA prepared the data, performed statistical analyses, together with VK and JD compiled the final neophyte lists, with MV prepared the maps and led the writing. All co-authors helped with species status assignment and commented on the manuscript.

The vegetation-plot data used in this study are stored in the European Vegetation Archive database (EVA; http://euroveg.org/eva-database) under the project number 56 â€" 2017-10-11. Maps can be downloaded from the Zenodo repository (link to be added).

Data availability statement

The vegetation-plot data used in this study are stored in the European Vegetation Archive database (EVA; http://euroveg.org/eva-database) under the project number 56 -2017-10-11. Maps can be downloaded from the Zenodo repository (link to be added).

Appendix S2 Overview of the habitat types. Appendix S3 List of regions. Appendix S4 Literature used for status assessment. Appendix S5 References used for life-form assessment of neophytes. Appendix S6 Summary statistics for Europe and regions. Appendix S7 Summary statistics for Europe and regions calculated across plots. Appendix S8 Summary statistics for habitats. Appendix S9 Summary statistics for habitats calculated across plots. Appendix S3 List of regions. Appendix S4 Literature used for status assessment. Appendix S5 References used for life-form assessment of neophytes. Appendix S6 Summary statistics for Europe and regions. Appendix S7 Summary statistics for Europe and regions calculated across plots. Appendix S8 Summary statistics for habitats. Appendix S9 Summary statistics for habitats calculated across plots. Appendix S10 List of neophytes recorded per region. Appendix S11 Comparison of relative neophyte species richness and neophyte cover among habitat groups. Appendix S12 Maps of the proportion of invaded plots in individual habitats relative to the neophyte origin categories.