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The goal of this task was to identify and characterise novel methods for biodiversity monitoring, and to assess their suitability for large scale deployment across Europe. To address this goal we combined extensive literature searches with expert consultation, namely using a survey and through an online workshop. The outcome of our searches is summarised in a metadatabase, which includes 282 methods or method components, which have been classified according to EBV classes addressed, target taxa, and broad method type the method relates to. We then consulted experts within the EuropaBON network and beyond, on the advantages and challenges associated with each of these novel methods, as well as their technology readiness level. In combination, our approaches revealed a wealth of novel methods and a highly active research field, with extensive emerging innovation on several fronts. However, it also revealed high variability in technology readiness, with lack of validation being a prevalent hurdle yet to be overcome for many applications of these methods (i.e. for some taxa and in some environments). Moreover, the opportunities for expansion in observations created by these novel approaches open new challenges associated to the standardisation, integration and storage of biodiversity monitoring data. Finally, the expansion of observations should take a designed approach, in order to deliver on its potential to improve representation and resolution of biodiversity monitoring, and should aim to complement rather than replace human observations.

Aims: Classification of European bog vegetation (Oxycocco-Sphagnetea class); identification of diagnostic species for the class and vegetation subgroups (orders and alliances); development of an expert system for automatic classification of vegetation plots; and production of distribution maps of the Oxycocco-Sphagnetea class and its alliances. Location: Europe. Methods: A data set of vegetation-plot records was compiled to include various bog types over most of the European continent. An unsupervised classification (beta-flexible linkage method, Sørensen distance measure) and detrended correspondence analysis (DCA) ordination were applied. Formal definitions of syntaxa based on species presence and covers, and respecting the results of the unsupervised classification, were developed and included in a classification expert system. Results: The Oxycocco-Sphagnetea class, its two orders (Sphagno-Ericetalia tetralicis and Sphagnetalia medii) and seven compositionally distinct alliances were formally defined. In addition to the syntaxa included in EuroVegChecklist, three new alliances were distinguished: Rubo chamaemori-Dicranion elongati (subarctic polygon and palsa mires); Erico mackaianae-Sphagnion papillosi (blanket bogs of the northwestern Iberian Peninsula); and Sphagno baltici-Trichophorion cespitosi (boreal bog lawns). The latter alliance is newly described in this article. Conclusions: This first pan-European formalized classification of European bog vegetation partially followed the system presented in EuroVegChecklist, but suggested three additional alliances. One covers palsa and polygon mires, one covers Iberian bogs with endemics and one fills the syntaxonomical gap for lawn microhabitats in boreal bogs. A classification expert system has been developed, which allows assignment of vegetation plots to the types described.

Rising temperatures may endanger fragile ecosystems because their character and key species show different habitat affinities under different climates. This assumption has only been tested in limited geographical scales. In fens, one of the most endangered ecosystems in Europe, broader pH niches have been reported from cold areas and are expected for colder past periods. We used the largest European-scale vegetation database from fens to test the hypothesis that pH interacts with macroclimate temperature in forming realized niches of fen moss and vascular plant species. We calibrated the data set (29,885 plots after heterogeneity-constrained resampling) with temperature, using two macroclimate variables, and with the adjusted pH, a variable combining pH and calcium richness. We modelled temperature, pH and water level niches for one hundred species best characterizing European fens using generalized additive models and tested the interaction between pH and temperature. Fifty-five fen species showed a statistically significant interaction between pH and temperature (adj p ˂ .01). Forty-six of them (84%) showed a positive interaction manifested by a shift or restriction of their niche to higher pH in warmer locations. Nine vascular plants and no moss showed the opposite interaction. Mosses showed significantly greater interaction. We conclude that climate significantly modulates edaphic niches of fen plants, especially bryophytes. This result explains previously reported regional changes in realized pH niches, a current habitat-dependent decline of endangered taxa, and distribution changes in the past. A warmer climate makes growing seasons longer and warmer, increases productivity, and may lower the water level. These effects prolong the duration and intensity of interspecific competition, support highly competitive Sphagnum mosses, and, as such, force niches of specialized fen species towards narrower high-pH ranges. Recent anthropogenic landscape changes pose a severe threat to many fen species and call for mitigation measures to lower competition pressure in their refugia.

Long-term analyses of biodiversity data highlight a 'biodiversity conservation paradox': biological communities show substantial species turnover over the past century1,2, but changes in species richness are marginal1,3-5. Most studies, however, have focused only on the incidence of species, and have not considered changes in local abundance. Here we asked whether analysing changes in the cover of plant species could reveal previously unrecognized patterns of biodiversity change and provide insights into the underlying mechanisms. We compiled and analysed a dataset of 7,738 permanent and semi-permanent vegetation plots from Germany that were surveyed between 2 and 54 times from 1927 to 2020, in total comprising 1,794 species of vascular plants. We found that decrements in cover, averaged across all species and plots, occurred more often than increments; that the number of species that decreased in cover was higher than the number of species that increased; and that decrements were more equally distributed among losers than were gains among winners. Null model simulations confirmed that these trends do not emerge by chance, but are the consequence of species-specific negative effects of environmental changes. In the long run, these trends might result in substantial losses of species at both local and regional scales. Summarizing the changes by decade shows that the inequality in the mean change in species cover of losers and winners diverged as early as the 1960s. We conclude that changes in species cover in communities represent an important but understudied dimension of biodiversity change that should more routinely be considered in time-series analyses.

This is a dataset of Ellenberg-type indicator values for European vascular plant species described in this article: Tichý L., Axmanová I., Dengler J., Guarino R., Jansen F., Midolo G., Nobis M.P., Van Meerbeek K., Aćić S., Attorre F., Bergmeier E., Biurrun I., Bonari G., Bruelheide H., Campos J.A., Čarni A., Chiarucci A., Ćuk M., Ćušterevska M., Didukh Y., Dítě D., Dítě Z., Dziuba T., Fanelli G., Fernández-Pascual E., Garbolino E., Gavilán R.G., Gégout J.-C., Graf U., Güler B., Hájek M., Hennekens S.M., Jandt U., Jašková A., Jiménez-Alfaro B., Julve P., Kambach S., Karger D.N., Karrer G., Kavgacı A., Knollová I., Kuzemko A., Küzmič F., Landucci F., Lengyel A., Lenoir J., Marcenò C., Moeslund J.E., Novák P., Pérez-Haase A., Peterka T., Pielech R., Pignatti A., Rašomavičius V., Rūsiņa S., Saatkamp A., Šilc U., Škvorc Ž., Theurillat J.-P., Wohlgemuth T. & Chytrý M. (2023) Ellenberg-type indicator values for European vascular plant species. Journal of Vegetation Science, 34, e13168. https://doi.org/10.1111/jvs.13168 The dataset contains: 1) Harmonized Ellenberg-type indicator values for light, temperature, moisture, reaction, nutrients, and salinity for European vascular plants. 2) Original indicator values from 13 datasets of Ellenberg-type indicator values: - Germany (Ellenberg & Leuschner 2010) - Austria (Karrer 1992) - Cantabrian Range (Jiménez-Alfaro et al. 2021) - Czech Republic (Chytrý et al. 2018) - European mires (Hájek et al. 2020) - France (Julve 2015) - Great Britain (Hill et al. 2000) - Greece (South Aegean) (Böhling et al. 2002) - Hungary (Borhidi 1995) - Italy (Guarino & La Rosa 2019, modified) - Saline habitats (Dítě et al. 2023) - Switzerland and the Alps (Landolt et al. 2010) - Ukraine (Didukh 2011) 3) Species nomenclature is standardized according to the Euro+Med PlantBase (http://europlusmed.org)., The final version of the paper (accepted 8th Dec 2022; published 22nd Dec 2022). Difference from version 1.0: Minor changes in newly added species values calculated from co-occurrence with other species in the EVA database. They were corrected using straight-line calibration.

B.J.-A. was funded by the Marie Curie Clarín-COFUND program of the Principality of Asturias-EU (ACB17-26) and the Spanish Research Agency (AEI/10.13039/501100011033)., Testolin, R., Attorre, F., Borchardt, P., Brand, R.F., Bruelheide, H., Chytrý, M., De Sanctis, M., Dolezal, J., Finckh, M., Haider, S., Hemp, A., Jandt, U., Kessler, M., Korolyuk, A.Y., Lenoir, J., Makunina, N., Malanson, G.P., Montesinos-Tubée, D.B., Noroozi, J., Nowak, A., Peet, R.K., Peyre, G., Sabatini, F.M., Šibík, J., Sklenář, P., Sylvester, S.P., Vassilev, K., Virtanen, R., Willner, W., Wiser, S.K., Zibzeev, E.G., Jiménez-Alfaro, B.

International audience; Questions. What are the functional trade-offs of vascular plant species in global alpine ecosystems? How is functional variation related to vegetation zones, climatic groups and biogeographic realms? What is the relative contribution of macroclimate and evolutionary history in shaping the functional variation of alpine plant communities? Location. Global. Methods. We compiled a data set of alpine vegetation with 5,532 geo-referenced plots, 1,933 species and six plant functional traits. We used principal component analysis to quantify functional trade-offs among species and trait probability density to assess the functional dissimilarity of alpine vegetation in different vegetation zones, climatic groups and biogeographic realms. We used multiple regression on distance matrices to model community functional dissimilarity against environmental and phylogenetic dissimilarity, controlling for geographic distance. Results. The first two PCA axes explained 66% of the species’ functional variation and were related to the leaf and stem economic spectra, respectively. Trait probability density was largely independent of vegetation zone and macroclimate but differed across biogeographic realms. The same pattern emerged for both species pool and community levels. The effects of environmental and phylogenetic dissimilarities on community functional dissimilarity had similar magnitude, while the effect of geographic distance was negligible. Conclusions. Plant species in alpine areas reflect the global variation of plant function, but with a predominant role of resource use strategies. Current macroclimate exerts a limited effect on alpine vegetation, mostly acting at the community level in combination with evolutionary history. Global alpine vegetation is functionally unrelated to the vegetation zones in which it is embedded, exhibiting strong functional convergence across regions

Based on plant occurrence data covering all parts of Germany, we investigated changes in the distribution of 2136 plant species between 1960 and 2017. We analyzed 29 million occurrence records over an area of ~350,000 km2 on a 5 × 5 km grid using temporal and spatiotemporal models and accounting for sampling bias. Since the 1960s, more than 70% of investigated plant species showed declines in nationwide occurrence. Archaeophytes (species introduced before 1492) most strongly declined but also native plant species experienced severe declines. In contrast, neophytes (species introduced after 1492) increased in their nationwide occurrence but not homogeneously throughout the country. Our analysis suggests that the strongest declines in native species already happened in the 1960s-1980s, a time frame in which often few data exist. Increases in neophytic species were strongest in the 1990s and 2010s. Overall, the increase in neophytes did not compensate for the loss of other species, resulting in a decrease in mean grid cell species richness of -1.9% per decade. The decline in plant biodiversity is a widespread phenomenon occurring in different habitats and geographic regions. It is likely that this decline has major repercussions on ecosystem functioning and overall biodiversity, potentially with cascading effects across trophic levels. The approach used in this study is transferable to other large-scale trend analyses using heterogeneous occurrence data.

1. Niche filtering predicts that abundant species in communities have similar traits that are suitable for the environment. However, niche filtering can operate on distinct axes of trait variation in response to different ecological conditions. Here, we use a trait-based approach to infer niche filtering processes and (a) test if abundant and rare species in grassland communities are differently positioned along distinct axes of trait variation, (b) determine if these trait variation axes, as well as phylogenetic and functional similarities, drive species relative abundance (above-ground cover) within communities, and (c) explore whether these relationships vary across grassland types and macro-climatic gradients. 2. We analysed species abundance in vegetation plots from temperate grasslands in Central Europe as a function of species position along three axes of trait variation: the ‘Plant Size Spectrum’ (PSS), the ‘Leaf Economics Spectrum’ (LES) and the ‘Life span/Clonality Spectrum’ (LCS). We also used phylogenetic and functional similarities in the multi-dimensional trait space as predictors of species abundance. We compared our results among alpine, wet, mesic and dry grasslands and tested whether the effect of the predictors on species abundance was significant across macro-climatic gradients. 3. Compared to abundant species, rare species in grassland communities were more commonly annual and non-clonal, had lower stature and smaller leaves and seeds, and relied on more acquisitive leaf economics. Our predictors significantly explained species abundance in approximately one-third of the plots. LES was the most important predictor across all plots, with the most prominent effect in alpine and dry grasslands and areas with more extreme temperatures. In contrast, in mesic and wet grasslands and grasslands located in warmer and less seasonal regions, species abundance was best predicted by phylogenetic similarities between species, with Poaceae species becoming more abundant. 4. Our study explored trait–abundance relationships for different community types across a large area and broad macro-climatic gradients. We conclude that niche filtering, and particularly resource-acquisition trade-offs, drives species abundance in temperate grassland communities of Central Europe. Our findings emphasize the interaction between local environmental conditions and plant function in determining community assembly.

International audience; Species turnover is ubiquitous. However, it remains unknown whether certain types of species are consistently gained or lost across different habitats. Here, we analysed the trajectories of 1827 plant species over time intervals of up to 78 years at 141 sites across mountain summits, forests, and lowland grasslands in Europe. We found, albeit with relatively small effect sizes, displacements of smaller- by larger-ranged species across habitats. Communities shifted in parallel towards more nutrient-demanding species, with species from nutrient-rich habitats having larger ranges. Because these species are typically strong competitors, declines of smaller-ranged species could reflect not only abiotic drivers of global change, but also biotic pressure from increased competition. The ubiquitous component of turnover based on species range size we found here may partially reconcile findings of no net loss in local diversity with global species loss, and link community-scale turnover to macroecological processes such as biotic homogenisation.

Datos disponibles en https://github.com/fmsabatini/sPlotOpen_Code, EU H2020 project BACI, Grant No. 640176 (...), Sabatini, F.M., Lenoir, J., Hattab, T., Arnst, E.A., Chytrý, M., Dengler, J., De Ruffray, P., Hennekens, S.M., Jandt, U., Jansen, F., Jiménez-Alfaro, B., Kattge, J., Levesley, A., Pillar, V.D., Purschke, O., Sandel, B., Sultana, F., Aavik, T., Aćić, S., Acosta, A.T.R., Agrillo, E., Alvarez, M., Apostolova, I., Arfin Khan, M.A.S., Arroyo, L., Attorre, F., Aubin, I., Banerjee, A., Bauters, M., Bergeron, Y., Bergmeier, E., Biurrun, I., Bjorkman, A.D., Bonari, G., Bondareva, V., Brunet, J., Čarni, A., Casella, L., Cayuela, L., Černý, T., Chepinoga, V., Csiky, J., Ćušterevska, R., De Bie, E., de Gasper, A.L., De Sanctis, M., Dimopoulos, P., Dolezal, J., Dziuba, T., El-Sheikh, M.A.E.-R.M., Enquist, B., Ewald, J., Fazayeli, F., Field, R., Finckh, M., Gachet, S., Galán-de-Mera, A., Garbolino, E., Gholizadeh, H., Giorgis, M., Golub, V., Alsos, I.G., Grytnes, J.-A., Guerin, G.R., Gutiérrez, A.G., Haider, S., Hatim, M.Z., Hérault, B., Hinojos Mendoza, G., Hölzel, N., Homeier, J., Hubau, W., Indreica, A., Janssen, J.A.M., Jedrzejek, B., Jentsch, A., Jürgens, N., Kącki, Z., Kapfer, J., Karger, D.N., Kavgacı, A., Kearsley, E., Kessler, M., Khanina, L., Killeen, T., Korolyuk, A., Kreft, H., Kühl, H.S., Kuzemko, A., Landucci, F., Lengyel, A., Lens, F., Lingner, D.V., Liu, H., Lysenko, T., Mahecha, M.D., Marcenò, C., Martynenko, V., Moeslund, J.E., Monteagudo Mendoza, A., Mucina, L., Müller, J.V., Munzinger, J., Naqinezhad, A., Noroozi, J., Nowak, A., Onyshchenko, V., Overbeck, G.E., Pärtel, M., Pauchard, A., Peet, R.K., Peñuelas, J., Pérez-Haase, A., Peterka, T., Petřík, P., Peyre, G., Phillips, O.L., Prokhorov, V., Rašomavičius, V., Revermann, R., Rivas-Torres, G., Rodwell, J.S., Ruprecht, E., Rūsiņa, S., Samimi, C., Schmidt, M., Schrodt, F., Shan, H., Shirokikh, P., Šibík, J., Šilc, U., Sklenář, P., Škvorc, Ž., Sparrow, B., Sperandii, M.G., Stančić, Z., Svenning, J.-C., Tang, Z., Tang, C.Q., Tsiripidis, I., Vanselow, K.A., Vásquez Martínez, R., Vassilev, K., Vélez-Martin, E., Venanzoni, R., Vibrans, A.C., Violle, C., Virtanen, R., von Wehrden, H., Wagner, V., Walker, D.A., Waller, D.M., Wang, H.-F., Wesche, K., Whitfeld, T.J.S., Willner, W., Wiser, S.K., Wohlgemuth, T., Yamalov, S., Zobel, M., Bruelheide, H.

In the rhizosphere, plants are exposed to a multitude of different biotic and abiotic factors, to which they respond by exuding a wide range of secondary root metabolites. So far, it has been unknown to which degree root exudate composition is species‐specific and is affected by land use, the local impact and local neighborhood under field conditions. In this study, root exudates of 10 common grassland species were analyzed, each five of forbs and grasses, in the German Biodiversity Exploratories using a combined phytometer and untargeted liquid chromatography‐mass spectrometry (LC‐MS) approach. Redundancy analysis and hierarchical clustering revealed a large set of semi‐polar metabolites common to all species in addition to species‐specific metabolites. Chemical richness and exudate composition revealed that forbs, such as Plantago lanceolata and Galium species, exuded more species‐specific metabolites than grasses. Grasses instead were primarily affected by environmental conditions. In both forbs and grasses, plant functional traits had only a minor impact on plant root exudation patterns. Overall, our results demonstrate the feasibility of obtaining and untargeted profiling of semi‐polar metabolites under field condition and allow a deeper view in the exudation of plants in a natural grassland community.

Climate change and nitrogen (N) deposition are among the most important drivers of biodiversity change at a global scale. Although commonly considered separately in applied research, conservation, and policy development, both empirical evidence and modelling studies indicate that these two stressors may act synergistically and trigger a more pronounced biodiversity loss than each of them alone. Interactive effects thus need to be considered in biodiversity conservation. In this study, we propose a new method for jointly assessing risk from both stressors to habitats by combining climate envelope modelling and the evaluation of exceedance of empirical critical loads for eutrophication through N deposition. Based on this approach we can calculate a combined risk metric, which is the scaled exceedance of the N critical load and climatic thresholds in the three-dimensional space of N deposition, temperature, and water limitation. We use a habitat map to project this risk metric for 34 EUNIS habitat types across Austria. Resulting risk maps indicated that about 16 % of the area of these natural and semi-natural habitat types is currently at risk from either N exceedance, drought or warming. Using combinations of future climatic (RCP2.5 and RCP8.5) and N deposition (business as usual, current legislation and maximum feasible reduction) scenarios, we predict that the area at risk might increase to up to 19% until 2050, with risk from N-deposition decreasing and risk from climate change increasing. Combined impacts of both stressors affect only up to 2% of the entire evaluated area, but are much more frequent in particular habitat types such as oligotrophic bogs and subalpine to alpine grasslands. We conclude that this method provides a useful screening procedure to identify and compare areas and habitats under combined risk from both stressors and to thus support prioritization and decision making in biodiversity conservation.

Understanding drivers of success for alien species can inform on potential future invasions. Recent conceptual advances highlight that species may achieve invasiveness via performance along at least three distinct dimensions: 1) local abundance, 2) geographic range size, and 3) habitat breadth in naturalized distributions. Associations among these dimensions and the factors that determine success in each have yet to be assessed at large geographic scales. Here, we combine data from over one million vegetation plots covering the extent of Europe and its habitat diversity with databases on species' distributions, traits, and historical origins to provide a comprehensive assessment of invasiveness dimensions for the European alien seed plant flora. Invasiveness dimensions are linked in alien distributions, leading to a continuum from overall poor invaders to super invaders - abundant, widespread aliens that invade diverse habitats. This pattern echoes relationships among analogous dimensions measured for native European species. Success along invasiveness dimensions was associated with details of alien species' introduction histories: earlier introduction dates were positively associated with all three dimensions, and consistent with theory-based expectations, species originating from other continents, particularly acquisitive growth strategists, were among the most successful invaders in Europe. Despite general correlations among invasiveness dimensions, we identified habitats and traits associated with atypical patterns of success in only one or two dimensions - for example, the role of disturbed habitats in facilitating widespread specialists. We conclude that considering invasiveness within a multidimensional framework can provide insights into invasion processes while also informing general understanding of the dynamics of species distributions., Deutsche Forschungsgemeinschaft (264740629) Grantová Agentura České Republiky (19-28491X) Grantová Agentura České Republiky (19-28807X) Grantová Agentura České Republiky (RVO 67985939) Austrian Science Fund (I 2086 - B29) Bundesministerium für Bildung und Forschung (01LC1807A) Eusko Jaurlaritza (IT299-10) National Research Foundation of Korea (2018R1C1B6005351) University of Latvia (AAp2016/B041//Zd2016/AZ03) Villum Fonden (16549)

Aims: Biodiversity is traditionally studied mostly at the species level, but biogeographical and macroecological studies at higher taxonomic levels can provide valuable insights into the evolutionary processes at large spatial scales. Our aim was to assess the representation of vascular plant families within different vegetation formations across Europe. Location: Europe. Methods: We used a data set of 816,005 vegetation plots from the European Vegetation Archive (EVA). For each plot, we calculated the relative species richness of each plant family as the number of species belonging to that family divided by the total number of species. We mapped the relative species richness, averaged across all plots in 50 km × 50 km grid cells, for each family and broad habitat groups: forests, grasslands, scrub and wetlands. We also calculated the absolute species richness and the Shannon diversity index for each family. Results: We produced 522 maps of mean relative species richness for a total of 152 vascular plant families occurring in forests, grasslands, scrub and wetlands. We found distinct spatial patterns for many combinations of families and habitat groups. The resulting series of 522 maps is freely available, both as images and GIS layers. Conclusions: The distinct spatial patterns revealed in the maps suggest that the relative species richness of plant families at the community level reflects the evolutionary history of individual families. We believe that the maps and associated data can inspire further biogeographical and macroecological studies and strengthen the ongoing integration of phylogenetic, functional and taxonomic diversity concepts., MV, IA, JPC, ZL, IK, AJ and MC were funded by the Czech Science Foundation, programme EXPRO (project no. 19-28491X); JDi by the Czech Science Foundation (18-02773S); IB and JAC by the Basque Government (IT936-16); AČ by the Slovenian Research Agency (ARRS, P1-0236); AK by the National Research Foundation of Ukraine (project no. 2020.01/0140); JŠ by the Slovak Research and Development Agency (APVV 16-0431); KV by the National Science Fund (Contract DCOST 01/7/19.10.2018).

Aim Plant functional traits summarize the main variability in plant form and function across taxa and biomes. We assess whether geographic range size, climatic niche size, and local abundance of plants can be predicted by sets of traits (trait syndromes) or are driven by single traits. Location Eurasia. Methods Species distribution maps were extracted from the Chorological Database Halle to derive information on the geographic range size and climatic niche size for 456 herbaceous, dwarf shrub and shrub species. We estimated local species abundances based on 740,113 vegetation plots from the European Vegetation Archive, where abundances were available as plant species cover per plot. We compiled a complete species-by-trait matrix of 20 plant functional traits from trait databases (TRY, BiolFlor and CLO-PLA). The relationships of species' geographic range size, climatic niche size and local abundance with single traits and trait syndromes were tested with multiple linear regression models. Results Generally, traits were more strongly related to local abundances than to broad-scale species distribution patterns in geographic and climatic space (range and niche size), but both were better predicted by trait combinations than by single traits. Local abundance increased with leaf area and specific leaf area (SLA). Geographic range size and climatic niche size both increased with SLA. While range size increased with plant height, niche size decreased with leaf carbon content. Conclusion Functional traits matter for species' abundance and distribution at both local and broad geographic scale. Local abundances are associated with different combinations of traits as compared to broad-scale distributions, pointing to filtering by different environmental and ecological factors acting at distinct spatial scales. However, traits related to the leaf economics spectrum were important for species' abundance and occurrence at both spatial scales. This finding emphasizes the general importance of resource acquisition strategies for the abundance and distribution of herbaceous, dwarf shrub and shrub species. Graduiertenforderung Sachsen-Anhalt (scholarship to MS), with additional support through institutional funds of Martin Luther University Halle-Wittenberg. MC and CM were funded by the Czech Science Foundation (project no. 19-28491X)

Aim European grassland communities are highly diverse, but patterns and drivers of their continental-scale diversity remain elusive. This study analyses taxonomic and functional richness in European grasslands along continental-scale temperature and precipitation gradients. Location Europe. Methods We quantified functional and taxonomic richness of 55,748 vegetation plots. Six plant traits, related to resource acquisition and conservation, were analysed to describe plant community functional composition. Using a null-model approach we derived functional richness effect sizes that indicate higher or lower diversity than expected given the taxonomic richness. We assessed the variation in absolute functional and taxonomic richness and in functional richness effect sizes along gradients of minimum temperature, temperature range, annual precipitation, and precipitation seasonality using a multiple general additive modelling approach. Results Functional and taxonomic richness was high at intermediate minimum temperatures and wide temperature ranges. Functional and taxonomic richness was low in correspondence with low minimum temperatures or narrow temperature ranges. Functional richness increased and taxonomic richness decreased at higher minimum temperatures and wide annual temperature ranges. Both functional and taxonomic richness decreased with increasing precipitation seasonality and showed a small increase at intermediate annual precipitation. Overall, effect sizes of functional richness were small. However, effect sizes indicated trait divergence at extremely low minimum temperatures and at low annual precipitation with extreme precipitation seasonality. Conclusions Functional and taxonomic richness of European grassland communities vary considerably over temperature and precipitation gradients. Overall, they follow similar patterns over the climate gradients, except at high minimum temperatures and wide temperature ranges, where functional richness increases and taxonomic richness decreases. This contrasting pattern may trigger new ideas for studies that target specific hypotheses focused on community assembly processes. And though effect sizes were small, they indicate that it may be important to consider climate seasonality in plant diversity studies. CCFB, MAJH and SH were supported by the ERC project (CoG SIZE 647224), IB by the Basque Government (IT936-16), MC by the Czech Science Foundation (19-28491X), and SR by the University of Latvia grant (AAp2016/B041//Zd2016/AZ03)

Pouteau, R., et al., [Aim] The number of naturalized (i.e. established) alien species has increased rapidly over recent centuries. Given the differences in environmental tolerances among species, little is known about what factors determine the extent to which the observed size of the naturalized range of a species and hence the extent to which the observed richness of naturalized species of a region approach their full potential. Here, we asked which region- and species-specific characteristics explain differences between observed and expected naturalizations. [Location] Global. [Time period] Present. [Major taxa studied] Vascular plants. [Methods] We determined the observed naturalized distribution outside Europe for 1,485 species endemic to Europe using the Global Naturalized Alien Flora (GloNAF) database and their expected distributions outside Europe using species distribution models. First, we investigated which of seven socio-economic factors related to introduction pathways, anthropogenic pressures and inventory effort best explained the differences between observed and expected naturalized European floras. Second, we examined whether distributional features, economic use and functional traits explain the extent to which species have filled their expected ranges outside Europe. Results: In terms of suitable area, more than 95% of expected naturalizations of European plants were not yet observed. Species were naturalized in only 4.2% of their suitable regions outside of Europe (range filling) and in 0.4% of their unsuitable regions (range expansion). Anthropogenic habitat disturbance primarily explained the difference between observed and expected naturalized European floras, as did the number of treaties relevant to invasive species. Species of ornamental and economic value and with large specific leaf area performed better at filling and expanding beyond their expected range. Main conclusions: The naturalization of alien plant species is explained by climate matching but also by the regional level of human development, the introduction pressure associated with the ornamental and economic values of the species and their adaptation to disturbed environments., Robin Pouteau was supported by the National Natural Science Foundation of China (grant number 31901176) and Taizhou University (2018YQ001). Carsten Meyer acknowledges funding by the Volkswagen Foundation through a Freigeist Fellowship (A118199), and additional support by iDiv, funded by the German Research Foundation (DFG–FZT 118, 202548816). Petr Pyšek and Jan Pergl were supported by the Czech Science Foundation (EXPRO grant 19-28807X) and the Czech Academy of Sciences (long-term research development project RVO 67985939). Idoia Biurrun and Juan Antonio Campos were supported by the Basque Government (IT936-16). Gianmaria Bonari, Milan Chytrý, Ilona Knollová, Anna Kuzemko and Corrado Marcenò were supported by the Czech Science Foundation (19-28491X). Rosario G. Gavilán was supported by Madrid Government project REMEDINAL (P2018/EMT-4338). Jens-Christian Svenning considers this work a contribution to his VILLUM Investigator project ‘Biodiversity Dynamics in a Changing World’ funded by VILLUM FONDEN (16549). Mark van Kleunen was supported by the German Research Foundation DFG (264740629). The authors appreciate the helpful comments by three anonymous referees and the Handling Editor, Melodie A. McGeoch.

Aim: This work explores whether the commonly observed positive range size-niche breadth relationship exists for Fagus, one of the most dominant and widespread broad-leaved deciduous tree genera in temperate forests of the Northern Hemisphere. Additionally, we ask whether the 10 extant Fagus species' niche breadths and climatic tolerances are under phylogenetic control. Location: Northern Hemisphere temperate forests. Taxon: Fagus L. Methods: Combining the global vegetation database sPlot with Chinese vegetation data, we extracted 107,758 releves containing Fagus species. We estimated biotic and climatic niche breadths per species using plot-based co-occurrence data and a resource-based approach, respectively. We examined the relationships of these estimates with range size and tested for their phylogenetic signal, prior to which a Random Forest (RF) analysis was applied to test which climatic properties are most conserved across the Fagus species. Results: Neither biotic niche breadth nor climatic niche breadth was correlated with range size, and the two niche breadths were incongruent as well. Notably, the widespread North American F. grandifolia had a distinctly smaller biotic niche breadth than the Chinese Fagus species (F. engleriana, F. hayatae, F. longipetiolata and F. lucida) with restricted distributions in isolated mountains. The RF analysis revealed that cold tolerance did not differ among the 10 species, and thus may represent an ancestral, fixed trait. In addition, neither biotic nor climatic niche breadths are under phylogenetic control. Main Conclusions: We interpret the lack of a general positive range size-niche breadth relationship within the genus Fagus as a result of the widespread distribution, high among-region variation in available niche space, landscape heterogeneity and Quaternary history. The results hold when estimating niche sizes either by fine-scale co-occurrence data or coarse-scale climate data, suggesting a mechanistic link between factors operating across spatial scales. Besides, there was no evidence for diverging ecological specialization within the genus Fagus. National Natural Science Foundation of ChinaNational Natural Science Foundation of China (NSFC); Ministry of Science and Technology of ChinaMinistry of Science and Technology, China; National Key Research and Development Program of China; Grantova Agentura Ceske RepublikyGrant Agency of the Czech Republic; Chinese Scholarship CouncilChina Scholarship Council; Independent Research Fund Denmark; Villum Fonden; German Research FoundationGerman Research Foundation (DFG); Ministry of Education, Youth and Sport of the Czech Republic, program Inter-Excellence; Natural Sciences project TREECHANGE National Natural Science Foundation of China; Ministry of Science and Technology of China; National Key Research and Development Program of China; Grantova Agentura Ceske Republiky; Chinese Scholarship Council; Independent Research Fund Denmark; Natural Sciences project TREECHANGE; Villum Fonden; German Research Foundation; Ministry of Education, Youth and Sport of the Czech Republic, program Inter-Excellence

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

Aim: To identify functional traits that best predict community assembly without knowing the underlying environmental drivers. Methods: We propose a new method based on the correlation r(XY) between two matrices of potential community composition: the matrix X is fuzzy-weighted by trait similarities of species, and the matrix Y is derived by Beals smoothing using the probabilities of species co-occurrences. Since X is based on one or more traits, r(XY) measures how well the traits used for fuzzy-weighting reflect the species co-occurrence patterns in Y. We developed an optimisation algorithm to identify the traits maximising this correlation, together with an appropriate permutational test for significance. Using metacommunity data generated by a stochastic, individual-based, spatially explicit model, we assessed the type I error and the power of our method across different simulation scenarios, varying environmental filtering parameters, number of traits and trait correlation structures. Then, we applied the method to real-world community and trait data of dry calcareous grassland communities across Germany to identify, out of 49 traits, the combination of traits that maximised r(XY). Results: The method correctly identified the relevant traits involved in the assembly mechanisms of simulated communities, showing high power and accurate type I error. It proved to be robust against confounding aspects related to interactions between environmental factors, strength of limiting factors, and trait collinearity. In the grassland dataset, the method identified five traits that best explained community assembly. These traits reflect the size and the leaf economics spectrum, which are related to succession and resource supply, factors that may not be always measured in real-world situations. Conclusions: Our method successfully identified the relevant traits mediating community assembly, therefore providing insights on the underlying environmental and biotic factors, even if these are hidden, unmeasured or not accessible at the spatial or temporal scale of the study.

[Questions] European woodlands harbor at least 386 alien plant species but the factors driving local invasions remain unknown. By using a large vegetation-plot database, we asked how local richness and abundance of alien species vary by regions, elevation, climate, soil properties, human disturbance, and habitat types., [Location] Western, central and southern Europe., [Methods] We linked consolidated data from the European Vegetation Archive (16,211 plots) to a habitat classification scheme, climate, soil properties and human disturbance variables. In addition, we used 250 km × 250 km regional grid cells to test whether local patterns differ among regions. We used generalized additive models (GAMs) and quantile GAMs to explore how relative alien species richness and the sum of alien species covers per plot relate to predictors. Random Forest analyses (RFs) were employed to assess the importance of individual predictors that were not multicollinear., [Results] Relative alien species richness and the sum of alien species covers varied across regions and habitat types, with effects being more pronounced at the maximum rather than average responses. Both response variables declined with increasing elevation and distance to the nearest road or railroad and increased with the amount of sealed soil. Maxima in fitted functions matched plots from regional invasion hotspots in northwestern and central Europe. RFs accounted for 39.6% and 20.9% of the total variation in relative alien species richness and the sum of alien species covers, respectively, with region and habitat being the most important variables., [Conclusions] The importance of maximum response quantiles and the prevalence of regional hotspots point to invasion debt in European woodlands. As alien plants expand further, their species richness and abundance in woodlands will be likely driven by the shared effects of the introduction and planting history, differences in the invaded habitat types, and dispersal corridors., VW initiated this study as a part of the InvasEVe project financed by the SoMoPro II program and considers it as part of her Discovery Grant from the Canadian Natural Science and Engineering Research Council. VW's research leading to these results has acquired a financial grant from the People Program (Marie Curie Action) of the Seventh Framework Program of the EU according to REA Grant Agreement No. 291782, and it was further co-financed by the South-Moravian Region. MV and MC were supported by the Czech Science Foundation (19-28491X). PP and JP were supported by the Czech Science Foundation (19-28807X) and the Czech Academy of Sciences (RVO 67985939). JCS considers this work a contribution to his VILLUM Investigator project “Biodiversity Dynamics in a Changing World” funded by VILLUM FONDEN (grant 16549). IB and JAC were funded by the Basque Government (IT936-16). FFG was funded by the Plan Propio of the UCLM (2020-GRIN-29214)

Im vorliegenden Beitrag identifizieren wir Möglichkeiten und Grenzen einer Aufarbeitung bisher noch ungenutzter Datenquellen für raum-zeitliche Biodiversitätsanalysen in Deutschland. Im Rahmen des Syntheseprojekts sMon (https://www.idiv.de/smon) des Deutschen Zentrums für integrative Biodiversitätsforschung (iDiv) Halle-Jena-Leipzig erprobt eine Arbeitsgemeinschaft aus Bundes- und Landesbehörden gemeinsam mit Fachgesellschaften, Verbänden und wissenschaftlichen Einrichtungen Methoden, wie heterogene Daten der letzten Jahrzehnte für retrospektive Trendanalysen genutzt werden können. Darauf aufbauend schlagen wir gangbare Wege zur Zukunft des deutschen Biodiversitätsmonitorings vor und illustrieren, wie Daten aus der Vergangenheit helfen können, Monitoring künftig zu gestalten.

Aim The EUNIS Habitat Classification is a widely used reference framework for European habitat types (habitats), but it lacks formal definitions of individual habitats that would enable their unequivocal identification. Our goal was to develop a tool for assigning vegetation-plot records to the habitats of the EUNIS system, use it to classify a European vegetation-plot database, and compile statistically-derived characteristic species combinations and distribution maps for these habitats. Location Europe. Methods We developed the classification expert system EUNIS-ESy, which contains definitions of individual EUNIS habitats based on their species composition and geographic location. Each habitat was formally defined as a formula in a computer language combining algebraic and set-theoretic concepts with formal logical operators. We applied this expert system to classify 1,261,373 vegetation plots from the European Vegetation Archive (EVA) and other databases. Then we determined diagnostic, constant and dominant species for each habitat by calculating species-to-habitat fidelity and constancy (occurrence frequency) in the classified data set. Finally, we mapped the plot locations for each habitat. Results Formal definitions were developed for 199 habitats at Level 3 of the EUNIS hierarchy, including 25 coastal, 18 wetland, 55 grassland, 43 shrubland, 46 forest and 12 man-made habitats. The expert system classified 1,125,121 vegetation plots to these habitat groups and 73,188 to other habitats, while 63,064 plots remained unclassified or were classified to more than one habitat. Data on each habitat were summarized in factsheets containing habitat description, distribution map, corresponding syntaxa and characteristic species combination. Conclusions EUNIS habitats were characterized for the first time in terms of their species composition and distribution, based on a classification of a European database of vegetation plots using the newly developed electronic expert system EUNIS-ESy. The data provided and the expert system have considerable potential for future use in European nature conservation planning, monitoring and assessment. The previous versions of the expert system and related reports were produced within a contract from the European Environment Agency to Wageningen Environmental Research and Masaryk University. The opinions expressed are those of the contractor and do not represent the Agency's official position. EVA data management and preparation of this paper were supported by the Czech Science Foundation (project no. 19-28491X to MC, LT, IK, TP, CM, JDa, MH, PN, DZ, GB, AJ, AKu, ZL and DV). IB and JAC were supported by the Basque Government (project no. T936-16). TB, ET, and LK were supported by the Ministry of Science and Higher Education of the Russian Federation (TB and ET project no. AAAA-A18-118052590019-7; LK project no. AAAA-A19-119012490096-2).

AimTo identify functional traits that best predict community assembly without knowing the driving environmental factors.MethodsWe propose a new method that is based on the correlation r(XY) between two matrices of potential community composition: matrix X is fuzzy-weighted by trait similarities of species, and matrix Y is derived by Beals smoothing using the probabilities of species co-occurrences. Since matrix X is based on one or more traits, r(XY) measures how well the traits used for fuzzy-weighting reflect the observed co-occurrence patterns. We developed an optimization algorithm that identifies those traits that maximize this correlation, together with an appropriate permutational test for significance. Using metacommunity data generated by a stochastic, individual-based, spatially explicit model, we assessed the type I error and the power of our method across different simulation scenarios, varying environmental filtering parameters, number of traits and trait correlation structures. We then applied the method to real-world community and trait data of dry calcareous grassland communities across Germany to identify, out of 49 traits, the combination of traits that maximizes r(XY).ResultsThe method correctly identified the relevant traits involved in the community assembly mechanisms specified in simulations. It had high power and accurate type I error and was robust against confounding aspects related to interactions between environmental factors, strength of limiting factors, and correlation among traits. In the grassland dataset, the method identified five traits that best explained community assembly. These traits reflected the size and the leaf economics spectrum, which are related to succession and resource supply, factors that may not be always measured in real-world situations.ConclusionsOur method successfully identified the relevant traits mediating community assembly driven by environmental factors which may be hidden for not being measured or accessible at the spatial or temporal scale of the study.

Based on plant occurrence data covering all parts of Germany, we investigated changes in the distribution of 2146 plant species between 1960 and 2017. We analyzed 29 million occurrence records over an area of ∼350.000 km2 on a 5 × 5 km grid using temporal and spatio-temporal models and accounting for sampling bias. Since the 1960s, more than 70% of investigated plant species showed significant declines in nation-wide occurrence. Archaeophytes (species introduced before 1492) most strongly declined but also native plant species experienced severe declines. In contrast, neophytes (species introduced after 1492) increased in their nation-wide occurrence but not homogeneously throughout the country. Our analysis suggests that the strongest declines in native species already happened in the 1960s-80s, a time frame in which usually few data exist. Increases in neophytic species were strongest in the 1990s and 2010s. Overall, the increase in neophytes did not compensate for the loss of other species, resulting in a decrease in mean grid-cell species-richness of -1.9% per decade. The decline in plant biodiversity is a widespread phenomenon occurring in different habitats and geographic regions. It is likely that this decline has major repercussions on ecosystem functioning and overall biodiversity, potentially with cascading effects across trophic levels. The approach used in this study is transferable to large-scale trend analyses using heterogeneous occurrence data.

Netherlands Organisation for Scientific Research (NWO), Grant/Award Number: 019.171LW.023; German Research Foundation; German Research Foundation DFG, Grant/Award Number: 264740629; FONDECYT, Grant/Award Number: 11150835; CONICYT-PAI, Grant/Award Number: 82130046; 7th European Community Framework Programme, Grant/Award Number: FORECOFUN-SSA PIEF-GA-2010–274798; European Research Council Synergy, Grant/Award Number: ERC-SyG-2013-610028; Austrian Science Foundation, Grant/Award Number: I2086-B16; Czech Science Foundation, Centre of Excellence PLADIAS, Grant/Award Number: 14-15414S and RVO 67985939; EXPRO, Grant/Award Number: 19-28807X; VEGA, Grant/Award Number: 0051/1, van der Sande, M.T., Bruelheide, H., Dawson, W., Dengler, J., Essl, F., Field, R., Haider, S., van Kleunen, M., Kreft, H., Pagel, J., Pergl, J., Purschke, O., Pyšek, P., Weigelt, P., Winter, M., Attorre, F., Aubin, I., Bergmeier, E., Chytrý, M., Dainese, M., De Sanctis, M., Fagundez, J., Golub, V., Guerin, G.R., Gutiérrez, A.G., Jandt, U., Jansen, F., Jiménez-Alfaro, B., Kattge, J., Kearsley, E., Klotz, S., Kramer, K., Moretti, M., Niinemets, Ü., Peet, R.K., Penuelas, J., Petřík, P., Reich, P.B., Sandel, B., Schmidt, M., Sibikova, M., Violle, C., Whitfeld, T.J.S., Wohlgemuth, T., Knight, T.M.

Aim: The loss of biodiversity has raised serious concerns about the entailing losses of ecosystem services. Here, we explore the potential of repeated habitat mapping data to identify floristic changes over time. Using one German federal state as a case study, we assessed floristic changes between the 1980s and 2010s. These habitat data have great potential for analysis because of their high spatial coverage while also posing methodological challenges such as incomplete observation data. We developed a modelling approach that accounts for incomplete observations and explored the ability to detect temporal trends. Location: The Federal State of Schleswig‐Holstein (Germany) Methods: We compiled plant species lists from the earliest (1980s) and most recent (2010s) habitat mapping survey and aligned differing habitat definitions across mapping campaigns. A total of 5,503 mapped polygons, each with a list of species records, intersected the two surveys. We accounted for underrecorded species by assigning occurrence probabilities, based on species co‐occurrence information across all surveys, using Beals' index and tested the robustness of this approach by simulation experiments. For those species with significant increases and decreases in occurrence probability, we linked these trends to the species' functional characteristics. Results: We found a systematic loss of species that are moderately threatened. Species that indicate low nitrogen supply and high soil moisture declined, suggesting a shift towards a more eutrophic and drier landscape. Importantly, assessing specific plant traits associated with losses, we also detected a decrease in species with reddish and blueish flowers and species providing nectar, pointing to a decrease of insect‐pollinated taxa. Main conclusions: The identified changes raise concerns that plant biodiversity has fundamentally changed over the last three decades, with concomitant consequences for ecosystem services, especially pollination. Given the general lack of historical standardized data, our approach for trend analyses using incomplete observation data may be widely applicable to assess long‐term biodiversity change.

Aim A fundamental question in macroecology centres around understanding the relationship between species' local abundance and their distribution in geographical and climatic space (i.e. the multi-dimensional climatic space or climatic niche). Here, we tested three macroecological hypotheses that link local abundance to the following range properties: (a) the abundance-range size relationship, (b) the abundance-range centre relationship and (c) the abundance-suitability relationship. Location Europe. Taxon Vascular plants. Methods Distribution range maps were extracted from the Chorological Database Halle to derive information on the range and niche sizes of 517 European vascular plant species. To estimate local abundance, we assessed samples from 744,513 vegetation plots in the European Vegetation Archive, where local species' abundance is available as plant cover per plot. We then calculated the 'centrality', that is, the distance between the location of the abundance observation and each species' range centre in geographical and climatic space. The climatic suitability of plot locations was estimated using coarse-grain species distribution models (SDMs). The relationships between centrality or climatic suitability with abundance was tested using linear models and quantile regression. We summarized the overall trend across species' regression slopes from linear models and quantile regression using a meta-analytical approach. Results We did not detect any positive relationships between a species' mean local abundance and the size of its geographical range or climatic niche. Contrasting yet significant correlations were detected between abundance and centrality or climatic suitability among species. Main conclusions Our results do not provide unequivocal support for any of the relationships tested, demonstrating that determining properties of species' distributions at large grains and extents might be of limited use for predicting local abundance, including current SDM approaches. We conclude that environmental factors influencing individual performance and local abundance are likely to differ from those factors driving plant species' distribution at coarse resolution and broad geographical extents. Graduiertenforderung Sachsen-Anhalt (scholarship to MS), with additional support through institutional funds of Martin Luther University Halle-Wittenberg. PK received support from the German Centre for Integrative Biodiversity Research (iDiv) Halle-Jena-Leipzig funded by the German Research Foundation (FZT 118). IB and JAC were supported by the Basque Government (IT936-16). MC, AK and CM were supported by the Czech Science Foundation (19-28491X). SR was supported by the University of Latvia through grant funding (AAp2016/B041//Zd2016/AZ03). We acknowledge the financial support of the Open Access Publication Fund of the Martin Luther University Halle-Wittenberg.

Aims: To create a comprehensive, consistent and unequivocal phytosociological classification of European marsh vegetation of the class Phragmito-Magnocaricetea. Location: Europe. Methods: We applied the Cocktail method to a European data set of 249, 800 vegetation plots. We identified the main purposes and attributes on which to base the classification, defined assignment rules for vegetation plots, and prepared formal definitions for all the associations, alliances and orders of the class Phragmito-Magnocaricetea using formal logic. Each formula consists of the combination of “functional species groups”, cover values of individual species, and in the case of high-rank syntaxa also of “discriminating species groups” created using the Group Improvement (GRIMP) method. Results: The European Phragmito-Magnocaricetea vegetation was classified into 92 associations grouped in 11 alliances and six orders. New syntaxa (previously invalidly published according to the International Code of Phytosociological Nomenclature) were introduced: Bolboschoeno maritimi-Schoenoplection tabernaemontani, Glycerio maximae-Sietum latifolii, Glycerio notatae-Veronicetum beccabungae, Schoenoplectetum corymbosi and Thelypterido palustris-Caricetum elongatae. Based on a critical revision, some other syntaxa were rejected or excluded from the class Phragmito-Magnocaricetea. Conclusions: This work provides the first consistent classification of the class Phragmito-Magnocaricetea at the European scale, which is an important tool for nature conservation. Our classification largely respects previously existing concepts of syntaxa, but it also proposes modifications to the recently published EuroVegChecklist. This work also provides a protocol that can be used for extending the current classification to new syntaxa and geographical regions.

Aim: To disentangle the influence of environmental factors at different spatial grains (regional and local) on fern and lycophyte species richness and ask how regional and plot-level richness are related to each other. Location: Global. Time period: Present. Major Taxa studied: Ferns and lycophytes. Methods: We explored fern and lycophyte species richness at two spatial grains, regional (hexagonal grid cells of 7,666 km2) and plot level (300–500 m2), in relation to environmental data at regional and local grains (the 7,666 km2 hexagonal grid cells and 4 km2 square grid cells, respectively). For the regional grain, we obtained species richness data for 1,243 spatial units and used them together with climatic and topographical predictors to model global fern richness. For the plot-level grain, we collated a global dataset of nearly 83,000 vegetation plots with a surface area in the range 300–500 m2 in which all fern and lycophyte species had been counted. We used structural equation modelling to identify which regional and local factors have the biggest effect on plot-level fern and lycophyte species richness worldwide. We investigate how plot-level richness is related to modelled regional richness at the plot's location. Results: Plot-level fern and lycophyte species richness were best explained by models allowing a link between regional environment and plot-level richness. A link between regional richness and plot-level richness was essential, as models without it were rejected, while models without the regional environment-plot-level richness link were still valid but had a worse goodness-of-fit value. Plot-level richness showed a hump-shaped relationship with regional richness. Main conclusions: Regional environment and regional fern and lycophyte species richness each are important determinants of plot-level richness, and the inclusion of one does not substitute the inclusion of the other. Plot-level richness increases with regional richness until a saturation point is reached, after which plot-level richness decreases despite increasing regional richness, possibly reflecting species interactions.

There is growing interest in the application of plant functional trait-based approaches for development of sustainable land-use strategies. In this context, one crucial task is to identify and measure plant traits, which respond to land-use intensity (response traits) and simultaneously have an impact on ecosystem functions (effect traits). We hypothesized that species-specific leaf chemical composition, which may function both as response and effect trait, can be derived from Attenuated Total Reflection Fourier Transform Infrared (ATR-FTIR) spectroscopy tools in combination with multivariate statistical methods We investigated leaf ATR-FTIR spectra of two grasses, Poa pratensis L. and Dactylis glomerata L., and one forb, Achillea millefolium L. collected in grassland plots along a land-use intensity gradient in three regions of Germany. ATR-FTIR spectra appear to function as biochemical fingerprints unique to each species. The spectral response to land-use intensity was not consistent among species and less apparent in the two grasses than in the forb species. Whereas land-use intensification enhanced protein and cellulose content in A. millefolium , giving rise to changes in six spectral bands in the frequency range of 1088–1699 cm −1 , only cellulose content increased in D. glomerata , affecting the bands of 1385–1394 cm −1 . Poa pratensis spectra exhibited minimal changes under the influence of land-use, only in the spectral bands of 1373–1375 cm −1 associated with suberin-like aliphatic compounds. Our findings suggest that some species’ leaf chemical composition is responsive to land-use intensity, and thus, may have a predictive value for ecosystem services provided by those species within grassland vegetation (i.e., herbage yield quality).

Plant functional traits are widely used to predict community productivity. However, they are rarely used to predict individual plant performance in grasslands. To assess the relative importance of traits compared to environment, we planted seedlings of 20 common grassland species as phytometers into existing grassland communities varying in land‐use intensity. After 1 year, we dug out the plants and assessed root, leaf, and aboveground biomass, to measure plant performance. Furthermore, we determined the functional traits of the phytometers and of all plants growing in their local neighborhood. Neighborhood impacts were analyzed by calculating community‐weighted means (CWM) and functional diversity (FD) of every measured trait. We used model selection to identify the most important predictors of individual plant performance, which included phytometer traits, environmental conditions (climate, soil conditions, and land‐use intensity), as well as CWM and FD of the local neighborhood. Using variance partitioning, we found that most variation in individual plant performance was explained by the traits of the individual phytometer plant, ranging between 19.30% and 44.73% for leaf and aboveground dry mass, respectively. Similarly, in a linear mixed effects model across all species, performance was best predicted by phytometer traits. Among all environmental variables, only including land‐use intensity improved model quality. The models were also improved by functional characteristics of the local neighborhood, such as CWM of leaf dry matter content, root calcium concentration, and root mass per volume as well as FD of leaf potassium and root magnesium concentration and shoot dry matter content. However, their relative effect sizes were much lower than those of the phytometer traits. Our study clearly showed that under realistic field conditions, the performance of an individual plant can be predicted satisfyingly by its functional traits, presumably because traits also capture most of environmental and neighborhood conditions.

QUESTIONS: To what extent is trait variation in grasses and forbs driven by land‐use intensity, climate, soil conditions and plant diversity of the local neighbourhood? Do grass and forb species differ in the degree of intraspecific trait variation? LOCATION: Managed grasslands in three regions of Germany. METHODS: Using a phytometer approach, we raised 20 common European grassland species (ten forbs and ten grasses) and planted them into 54 plots of different land‐use types (pasture, meadow, mown pasture). After 1 yr in the field, we measured above‐ and below‐ground plant functional traits. Linear mixed effects models (LMEM) were used to identify the most powerful predictors for every trait. Variation partitioning was applied to assess the amount of inter‐ and intraspecific trait variation in grasses and forbs explained by environmental conditions (land‐use intensity, climate and soil conditions) and plant species diversity of the local neighbourhood. RESULTS: For 12 out of the 14 traits studied, either land‐use intensity or local neighbourhood diversity were predictors in the best LMEM. Land‐use intensity had considerably stronger effects than neighbourhood diversity. Root dry matter content and root phosphorus concentration of forbs were more affected by land‐use intensity than those of grasses. For almost all traits, intraspecific trait variation of grasses was much higher than that of forbs, while traits of forbs varied more among species. Overall, inter‐ and intraspecific variation was of the same magnitude. CONCLUSION: The similar magnitude of intra‐ and interspecific trait variation suggests that both sources should be considered in grassland studies at a scale similar to that of our study. The high amount of intraspecific trait variation that was explained by environmental factors and local neighbourhood diversity clearly demonstrates the high potential of species to adjust to local conditions, which would be ignored when only considering species mean trait values.

The loss of biodiversity at the global scale has been difficult to reconcile with observations of no net loss at local scales. Vegetation surveys across European temperate forests show that this may be explained by the replacement of small-ranged species with large-ranged ones, driven by nitrogen deposition. Biodiversity time series reveal global losses and accelerated redistributions of species, but no net loss in local species richness. To better understand how these patterns are linked, we quantify how individual species trajectories scale up to diversity changes using data from 68 vegetation resurvey studies of seminatural forests in Europe. Herb-layer species with small geographic ranges are being replaced by more widely distributed species, and our results suggest that this is due less to species abundances than to species nitrogen niches. Nitrogen deposition accelerates the extinctions of small-ranged, nitrogen-efficient plants and colonization by broadly distributed, nitrogen-demanding plants (including non-natives). Despite no net change in species richness at the spatial scale of a study site, the losses of small-ranged species reduce biome-scale (gamma) diversity. These results provide one mechanism to explain the directional replacement of small-ranged species within sites and thus explain patterns of biodiversity change across spatial scales.

Aim Biodiversity databases are valuable resources for understanding plant species distributions and dynamics, but they may insufficiently represent the actual geographic distribution and climatic niches of species. Here we propose and test a method to assess sampling coverage of species distribution in biodiversity databases in geographic and climatic space. Location Europe. Methods Using a test selection of 808,794 vegetation plots from the European Vegetation Archive (EVA), we assessed the sampling coverage of 564 European vascular plant species across both their geographic ranges and realized climatic niches. Range maps from the Chorological Database Halle (CDH) were used as background reference data to capture species geographic ranges and to derive species climatic niches. To quantify sampling coverage, we developed a box-counting method, the Dynamic Match Coefficient (DMC), which quantifies how much a set of occurrences of a given species matches with its geographic range or climatic niche. DMC is the area under the curve measuring the match between occurrence data and background reference (geographic range or climatic niche) across grids with variable resolution. High DMC values indicate good sampling coverage. We applied null models to compare observed DMC values with expectations from random distributions across species ranges and niches. Results Comparisons with null models showed that, for most species, actual distributions within EVA are deviating from null model expectations and are more clumped than expected in both geographic and climatic space. Despite high interspecific variation, we found a positive relationship in DMC values between geographic and climatic space, but sampling coverage was in general more random across geographic space. Conclusion Because DMC values are species-specific and most biodiversity databases are clearly biased in terms of sampling coverage of species occurrences, we recommend using DMC values as covariates in macroecological models that use species as the observation unit. This article is protected by copyright. All rights reserved.

Plants exude a diverse cocktail of metabolites into the soil as response to exogenous and endogenous factors. So far, root exudates have mainly been studied under artificial conditions due to methodological difficulties. In this study, each five perennial grass and forb species were investigated for polar and semi-polar metabolites in exudates under field conditions. Metabolite collection and untargeted profiling approaches combined with a novel classification method allowed the designation of 182 metabolites. The composition of exuded polar metabolites depended mainly on the local environment, especially soil conditions, whereas the pattern of semi-polar metabolites was primarily affected by the species identity. The profiles of both polar and semi-polar metabolites differed between growth forms, with grass species being generally more similar to each other and more responsive to the abiotic environment than forb species. This study demonstrated the feasibility of investigating exudates under field conditions and to identify the driving factors of exudate composition.