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Modelling the distribution and compositional variation of plant communities at the continental scale
Jiménez-Alfaro, Borja ; Suárez-Seoane, Susana ; Chytrý, Milan ; Hennekens, Stephan M. ; Willner, Wolfgang ; Hájek, Michal ; Agrillo, Emiliano ; Álvarez-Martínez, Jose M. ; Bergamini, Ariel ; Brisse, Henry ; Brunet, Jörg ; Casella, Laura ; Dítě, Daniel ; Font, Xavier ; Gillet, François ; Hájková, Petra ; Jansen, Florian ; Jandt, Ute ; Kącki, Zygmunt ; Lenoir, Jonathan ; Rodwell, John S. ; Schaminée, Joop H.J. ; Sekulová, Lucia ; Šibík, Jozef ; Škvorc, Željko ; Tsiripidis, Ioannis - \ 2018
Diversity and Distributions 24 (2018)7. - ISSN 1366-9516 - p. 978 - 990.
community distribution models - ecosystem properties - extent of occurrence - generalized dissimilarity modelling - habitat conservation - plant communities - vegetation
Aim: We investigate whether (1) environmental predictors allow to delineate the distribution of discrete community types at the continental scale and (2) how data completeness influences model generalization in relation to the compositional variation of the modelled entities. Location: Europe. Methods: We used comprehensive datasets of two community types of conservation concern in Europe: acidophilous beech forests and base-rich fens. We computed community distribution models (CDMs) calibrated with environmental predictors to predict the occurrence of both community types, evaluating geographical transferability, interpolation and extrapolation under different scenarios of sampling bias. We used generalized dissimilarity modelling (GDM) to assess the role of geographical and environmental drivers in compositional variation within the predicted distributions. Results: For the two community types, CDMs computed for the whole study area provided good performance when evaluated by random cross-validation and external validation. Geographical transferability provided lower but relatively good performance, while model extrapolation performed poorly when compared with interpolation. Generalized dissimilarity modelling showed a predominant effect of geographical distance on compositional variation, complemented with the environmental predictors that also influenced habitat suitability. Main conclusions: Correlative approaches typically used for modelling the distribution of individual species are also useful for delineating the potential area of occupancy of community types at the continental scale, when using consistent definitions of the modelled entity and high data completeness. The combination of CDMs with GDM further improves the understanding of diversity patterns of plant communities, providing spatially explicit information for mapping vegetation diversity and related habitat types at large scales.
Classification of European and Mediterranean coastal dune vegetation
Marcenò, Corrado ; Guarino, Riccardo ; Loidi, Javier ; Herrera, Mercedes ; Isermann, Maike ; Knollová, Ilona ; Tichý, Lubomír ; Tzonev, Rossen T. ; Acosta, Alicia Teresa Rosario ; Fitzpatrick, Úna ; Iakushenko, Dmytro ; Janssen, John A.M. ; Jiménez-Alfaro, Borja ; Kacki, Zygmunt ; Keizer-Sedláková, Iva ; Kolomiychuk, Vitaliy ; Rodwell, John S. ; Schaminée, Joop H.J. ; Šilc, Urban ; Chytrý, Milan - \ 2018
Applied Vegetation Science 21 (2018)3. - ISSN 1402-2001 - p. 533 - 559.
Ammophiletea - Biogeography - Expert system - Honckenyo-Elymetea - Koelerio-Corynephoretea canescentis - Phytosociology - Sand dune - Vegetation classification

Aims: Although many phytosociological studies have provided detailed local and regional descriptions of coastal dune vegetation, a unified classification of this vegetation in Europe and the Mediterranean Basin has been missing. Our aim is to produce a formalized classification of this vegetation and to identify the main factors driving its plant species composition at a continental scale. Location: Atlantic and Baltic coasts of Europe, Mediterranean Basin and the Black Sea region. Methods: We compiled a database of 30,759 plots of coastal vegetation, which were resampled to reduce unbalanced sampling effort, obtaining a data set of 11,769 plots. We classified these plots with TWINSPAN, interpreted the resulting clusters and used them for developing formal definitions of phytosociological alliances of coastal dune vegetation, which were included in an expert system for automatic vegetation classification. We related the alliances to climatic factors and described their biogeographic features and their position in the coastal vegetation zonation. We examined and visualized the floristic relationships among these alliances by means of DCA ordination. Results: We defined 18 alliances of coastal dune vegetation, including the newly described Centaureo cuneifoliae-Verbascion pinnatifidi from the Aegean region. The main factors underlying the differentiation of these alliances were biogeographic and macroclimatic contrasts between the Atlantic-Baltic, Mediterranean and Black Sea regions, along with ecological differences between shifting and stable dunes. The main difference in species composition was between the Atlantic-Baltic and Mediterranean-Black Sea regions. Within the former region, the main difference was driven by the different ecological conditions between shifting and stable dunes, whereas within the latter, the main difference was biogeographic between the Mediterranean and the Black Sea. Conclusions: The first formal classification of the European coastal dune vegetation was established, accompanied by an expert system containing the formal definitions of alliances, which can be applied to new data sets. The new classification system critically revised the previous concepts and integrated them into a consistent framework, which reflects the main gradients in species composition driven by biogeographic influences, macroclimate and the position of the sites in the coast-inland zonation of the dune systems. A revision of the class concept used in EuroVegChecklist is also proposed.

Alien plant invasions in European woodlands
Wagner, Viktoria ; Chytrý, Milan ; Jiménez-Alfaro, Borja ; Pergl, Jan ; Hennekens, Stephan ; Biurrun, Idoia ; Knollová, Ilona ; Berg, Christian ; Vassilev, Kiril ; Rodwell, John S. ; Škvorc, Željko ; Jandt, Ute ; Ewald, Jörg ; Jansen, Florian ; Tsiripidis, Ioannis ; Botta-Dukát, Zoltán ; Casella, Laura ; Attorre, Fabio ; Rašomavičius, Valerijus ; Ćušterevska, Renata ; Schaminée, Joop H.J. ; Brunet, Jörg ; Lenoir, Jonathan ; Svenning, Jens Christian ; Kącki, Zygmunt ; Petrášová-Šibíková, Mária ; Šilc, Urban ; García-Mijangos, Itziar ; Campos, Juan Antonio ; Fernández-González, Federico ; Wohlgemuth, Thomas ; Onyshchenko, Viktor ; Pyšek, Petr - \ 2017
Diversity and Distributions 23 (2017)9. - ISSN 1366-9516 - p. 969 - 981.
EUNIS - exotic - forest - invasive plants - life-form - neophyte - non-native - origin - tree
Aim: Woodlands make up a third of European territory and carry out important ecosystem functions, yet a comprehensive overview of their invasion by alien plants has never been undertaken across this continent. Location: Europe. Methods: We extracted data from 251,740 vegetation plots stored in the recently compiled European Vegetation Archive. After filtering (resulting in 83,396 plots; 39 regions; 1970–2015 time period), we analysed the species pool and frequency of alien vascular plants with respect to geographic origin and life-forms, and the levels of invasion across the European Nature Information System (EUNIS) woodland habitats. Results: We found a total of 386 alien plant species (comprising 7% of all recorded vascular plants). Aliens originating from outside of and from within Europe were almost equally represented in the species pool (192 vs. 181 species) but relative frequency was skewed towards the former group (77% vs. 22%) due, to some extent, to the frequent occurrence of Impatiens parviflora (21% frequency among alien plants). Phanerophytes were the most species-rich life-form (148 species) and had the highest representation in terms of relative frequency (39%) among aliens in the dataset. Apart from Europe (181 species), North America was the most important source of alien plants (109 species). At the local scale, temperate and boreal softwood riparian woodland (5%) and mire and mountain coniferous woodland (<1%) had the highest and lowest mean relative alien species richness (percentage of alien species per plot), respectively. Main conclusions: Our results indicate that European woodlands are prone to alien plant invasions especially when exposed to disturbance, fragmentation, alien propagule pressure and high soil nutrient levels. Given the persistence of these factors in the landscape, competitive alien plant species with a broad niche, including alien trees and shrubs, are likely to persist and spread further into European woodlands.
Diversity of lowland hay meadows and pastures in Western and Central Europe
Rodríguez-Rojo, Maria Pilar ; Jiménez-Alfaro, Borja ; Jandt, Ute ; Bruelheide, Helge ; Rodwell, John S. ; Schaminée, Joop H.J. ; Perrin, Philip M. ; Kacki, Zygmunt ; Willner, Wolfgang ; Fernández-González, Federico ; Chytrý, Milan - \ 2017
Applied Vegetation Science 20 (2017)4. - ISSN 1402-2001 - p. 702 - 719.
Grassland - Grazing - Management - Meadow - Mowing - Pasture - Phytosociology - Relevé - Vegetation classification - Vegetation database - Vegetation plot
Questions: Which are the main vegetation types of lowland hay meadows and pastures in Western and Central Europe? What are the main environmental gradients that drive patterns of species composition? Is it possible to classify these grasslands to phytosociological alliances that reflect management practices? Location: Western and Central Europe (excluding the Alps and Carpathians). Methods: A database of 21 400 vegetation plots of mesic grasslands across Western and Central Europe was compiled. After geographically stratified resampling, semi-supervised classification based on the K-means algorithm was applied to assign a subset of plots into 32 a priori association-level vegetation types and to search for new types within the subset of non-assigned plots. The vegetation plots assigned into the final vegetation types were submitted to another K-means classification to show the grouping into higher-level vegetation types. Results: A total of 36 associations were distinguished in the resampled subset of 8277 vegetation plots and were grouped into four large groups: (1) eutrophic and intensively managed hay meadows and permanent pastures; (2) nutrient-rich grasslands developed from recently abandoned fields or managed under irregular practices of mowing and manuring; (3) non-eutrophic lowland and submontane hay meadows; (4) extensively managed pastures and Atlantic grazed hay meadows. A PCoA of the associations of these four groups showed that extensively managed pastures were floristically more similar to non-eutrophic hay meadows than to permanent intensively managed pastures, which was more obvious in the Atlantic region than in Central Europe. Species composition of the lowland hay meadows was clearly differentiated according to biogeographic sectors. Other floristic differences were related to climate, altitude, soil base status and topography. Conclusions: This analysis challenges the traditional concept of mesic grassland alliances separating hay meadows from pastures. New classification should be based mainly on the differences in management intensity rather than in management practice. Consequently, nutrient-poor extensive pastures, which currently are not considered in the European Habitats Directive, should receive the same conservation attention as low-intensive hay meadows, because both types of vegetation can be equally species-rich and do not differ substantially in floristic composition from each other.
Classification of European beech forests : a Gordian Knot?
Willner, Wolfgang ; Jiménez-Alfaro, Borja ; Agrillo, Emiliano ; Biurrun, Idoia ; Campos, Juan Antonio ; Čarni, Andraž ; Casella, Laura ; Csiky, János ; Ćušterevska, Renata ; Didukh, Yakiv P. ; Ewald, Jörg ; Jandt, Ute ; Jansen, Florian ; Kącki, Zygmunt ; Kavgacı, Ali ; Lenoir, Jonathan ; Marinšek, Aleksander ; Onyshchenko, Viktor ; Rodwell, John S. ; Schaminée, Joop H.J. ; Šibík, Jozef ; Škvorc, Željko ; Svenning, Jens Christian ; Tsiripidis, Ioannis ; Turtureanu, Pavel Dan ; Tzonev, Rossen ; Vassilev, Kiril ; Venanzoni, Roberto ; Wohlgemuth, Thomas ; Chytrý, Milan - \ 2017
Applied Vegetation Science 20 (2017)3. - ISSN 1402-2001 - p. 494 - 512.
Beech forest - Europe - Fagetalia sylvaticae - Fagion sylvaticae - Fagus sylvatica - Luzulo-Fagion sylvaticae - Syntaxonomy - TWINSPAN - Vegetation plot database

Questions: What are the main floristic patterns in European beech forests? Which classification at the alliance and suballiance level is the most convincing?. Location: Europe and Asia Minor. Methods: We applied a TWINSPAN classification to a data set of 24 605 relevés covering the whole range of Fagus sylvatica forests and the western part of Fagus orientalis forests. We identified 24 ‘operational phytosociological units’ (OPUs), which were used for further analysis. The position of each OPU along the soil pH and temperature gradient was evaluated using Ellenberg Indicator Values. Fidelity of species to OPUs was calculated using the phi coefficient and constancy ratio. We compared alternative alliance concepts, corresponding to groups of OPUs, in terms of number and frequency of diagnostic species. We also established formal definitions for the various alliance concepts based on comparison of the total cover of the diagnostic species groups, and evaluated alternative geographical subdivisions of beech forests. Results: The first and second division levels of TWINSPAN followed the temperature and soil pH gradients, while lower divisions were mainly geographical. We grouped the 22 OPUs of Fagus sylvatica forests into acidophytic, meso-basiphytic and thermo-basiphytic beech forests, and separated two OPUs of F. orientalis forests. However, a solution with only two ecologically defined alliances of F. sylvatica forests (acidophytic vs basiphytic) was clearly superior with regard to number and frequency of diagnostic species. In contrast, when comparing groupings with three to six geographical alliances of basiphytic beech forests, respectively, we did not find a strongly superior solution. Conclusions: We propose to classify F. sylvatica forests into 15 suballiances – three acidophytic and 12 basiphytic ones. Separating these two groups at alliance or order level was clearly supported by our results. Concerning the grouping of the 12 basiphytic suballiances into ecological or geographical alliances, as advocated by many authors, we failed to find an optimal solution. Therefore, we propose a multi-dimensional classification of basiphytic beech forests, including both ecological and geographical groups as equally valid concepts which may be used alternatively depending on the purpose and context of the classification.

Formalized classification of European fen vegetation at the alliance level
Peterka, Tomáš ; Hájek, Michal ; Jiroušek, Martin ; Jiménez-Alfaro, Borja ; Aunina, Liene ; Bergamini, Ariel ; Dítě, Daniel ; Felbaba-Klushyna, Ljuba ; Graf, Ulrich ; Hájková, Petra ; Hettenbergerová, Eva ; Ivchenko, Tatiana G. ; Jansen, Florian ; Koroleva, Natalia E. ; Lapshina, Elena D. ; Lazarević, Predrag M. ; Moen, Asbjørn ; Napreenko, Maxim G. ; Pawlikowski, Paweł ; Plesková, Zuzana ; Sekulová, Lucia ; Smagin, Viktor A. ; Tahvanainen, Teemu ; Thiele, Annett ; Biţǎ-Nicolae, Claudia ; Biurrun, Idoia ; Brisse, Henry ; Ćušterevska, Renata ; Bie, Els De; Ewald, Jörg ; FitzPatrick, Úna ; Font, Xavier ; Jandt, Ute ; Kącki, Zygmunt ; Kuzemko, Anna ; Landucci, Flavia ; Moeslund, Jesper E. ; Pérez-Haase, Aaron ; Rašomavičius, Valerijus ; Rodwell, John S. ; Schaminée, Joop H.J. ; Šilc, Urban ; Stančić, Zvjezdana ; Chytrý, Milan ; Schwabe-Kratochwil, Angelika - \ 2017
Applied Vegetation Science 20 (2017)1. - ISSN 1402-2001 - p. 124 - 142.
Biogeography - Ecological gradients - Endangered habitats - Mires - Relevés - Supervised vegetation classification - Unsupervised vegetation classification - Vegetation plots - Wetlands
Aims: Phytosociological classification of fen vegetation (Scheuchzerio palustris-Caricetea fuscae class) differs among European countries. Here we propose a unified vegetation classification of European fens at the alliance level, provide unequivocal assignment rules for individual vegetation plots, identify diagnostic species of fen alliances, and map their distribution. Location: Europe, western Siberia and SE Greenland. Methods: 29 049 vegetation-plot records of fens were selected from databases using a list of specialist fen species. Formal definitions of alliances were created using the presence, absence and abundance of Cocktail-based species groups and indicator species. DCA visualized the similarities among the alliances in an ordination space. The ISOPAM classification algorithm was applied to regional subsets with homogeneous plot size to check whether the classification based on formal definitions matches the results of unsupervised classifications. Results: The following alliances were defined: Caricion viridulo-trinervis (sub-halophytic Atlantic dune-slack fens), Caricion davallianae (temperate calcareous fens), Caricion atrofusco-saxatilis (arcto-alpine calcareous fens), Stygio-Caricion limosae (boreal topogenic brown-moss fens), Sphagno warnstorfii-Tomentypnion nitentis (Sphagnum-brown-moss rich fens), Saxifrago-Tomentypnion (continental to boreo-continental nitrogen-limited brown-moss rich fens), Narthecion scardici (alpine fens with Balkan endemics), Caricion stantis (arctic brown-moss rich fens), Anagallido tenellae-Juncion bulbosi (Ibero-Atlantic moderately rich fens), Drepanocladion exannulati (arcto-boreal-alpine non-calcareous fens), Caricion fuscae (temperate moderately rich fens), Sphagno-Caricion canescentis (poor fens) and Scheuchzerion palustris (dystrophic hollows). The main variation in the species composition of European fens reflected site chemistry (pH, mineral richness) and sorted the plots from calcareous and extremely rich fens, through rich and moderately rich fens, to poor fens and dystrophic hollows. ISOPAM classified regional subsets according to this gradient, supporting the ecological meaningfulness of this classification concept on both the regional and continental scale. Geographic/macroclimatic variation was reflected in the second most important gradient. Conclusions: The pan-European classification of fen vegetation was proposed and supported by the data for the first time. Formal definitions developed here allow consistent and unequivocal assignment of individual vegetation plots to fen alliances at the continental scale.
Review of grassland habitats and development of distribution maps of heathland, scrub and tundra habitats of EUNIS habitats classification
Schaminee, J.H.J. ; Chytrý, M. ; Hennekens, S.M. ; Janssen, J.A.M. ; Jimenez-Alfaro, B. ; Knollová, Ilona ; Marceno, L. ; Mucina, L. ; Rodwell, J.S. ; Tichý, L. - \ 2016
EEA - European Environment Agency (Report EEA/NSV 15/005) - 372 p.
European Vegetation Archive (EVA) : An integrated database of European vegetation plots
Chytrý, Milan ; Hennekens, S.M. ; Jiménez-Alfaro, Borja ; Schaminée, J.H.J. ; Haveman, Rense ; Janssen, J.A.M. - \ 2016
Applied Vegetation Science 19 (2016)1. - ISSN 1402-2001 - p. 173 - 180.
Biodiversity informatics - Database - Ecoinformatics - European Vegetation Survey - International Association for Vegetation Science - Phytosociological data - Relevé - Vegetation database - Vegetation plot

The European Vegetation Archive (EVA) is a centralized database of European vegetation plots developed by the IAVS Working Group European Vegetation Survey. It has been in development since 2012 and first made available for use in research projects in 2014. It stores copies of national and regional vegetation- plot databases on a single software platform. Data storage in EVA does not affect on-going independent development of the contributing databases, which remain the property of the data contributors. EVA uses a prototype of the database management software TURBOVEG 3 developed for joint management of multiple databases that use different species lists. This is facilitated by the SynBioSys Taxon Database, a system of taxon names and concepts used in the individual European databases and their corresponding names on a unified list of European flora. TURBOVEG 3 also includes procedures for handling data requests, selections and provisions according to the approved EVA Data Property and Governance Rules. By 30 June 2015, 61 databases from all European regions have joined EVA, contributing in total 1 027 376 vegetation plots, 82% of them with geographic coordinates, from 57 countries. EVA provides a unique data source for large-scale analyses of European vegetation diversity both for fundamental research and nature conservation applications. Updated information on EVA is available online at

European Vegetation Archive (EVA): a new integrated source of European vegetation-plot data
Chytry, M. ; Hennekens, S.M. ; Jiménez-Alfaro, B. ; Knollová, I. ; Dengler, J. ; Schaminee, J.H.J. - \ 2014
In: Book of Abstracts, Biodiversity & Vegetation. Patterns, Processes, Conservation. - - p. 81 - 82.
Broad-scale distribution modelling of community types: an example using European vegetation-plot databases and MaxEnt
Jiménez-Alfaro, B. ; Suárez-Seoan, S. ; Chytry, M. ; Hennekens, S.M. ; Schaminee, J.H.J. ; Rodwell, J.S. - \ 2014
In: Book of Abstracts, Biodiversity & Vegetation. Patterns, Processes, Conservation. - - p. 115 - 116.
European Vegetation Archive (EVA): a new integrated source of European vegetation-plot data
Chytry, M. ; Hennekens, S.M. ; Jiménez-Alfaro, B. ; Knollová, I. ; Dengler, J. ; Schaminee, J.H.J. - \ 2014
Biogeographic patterns of base-rich fen vegetation across Europe
Jiménez-Alfaro, B. ; Hájek, M. ; Ejrnaes, R. ; Rodwell, J. ; Pawlikowski, P. ; Weeda, E.J. - \ 2014
Applied Vegetation Science 17 (2014)2. - ISSN 1402-2001 - p. 367 - 380.
environmental gradients - ecological gradients - plant associations - vascular plants - temperate zone - plot size - north - classification - mires - water
Questions What is the distribution of base-rich fen vegetation and the specialist species along European biogeographic regions? How do the gradients in species composition correlate to geography and climate at continental scale? What are the implications of such patterns for the classification of these habitats? Location Fifteen countries of Central, Western and Northern Europe. Methods We compiled a vegetation plot database of base-rich fens and related communities including vascular plants and bryophytes. The initial data set with 6943 plots was filtered according to the presence of specialists using discriminant analysis. We used DCA to analyse the correlation of species composition with geography and climate, and kriging interpolation for mapping gradients in the study area. Modified TWINSPAN was used to detect major vegetation groups. The results of the whole data set (plot size 1–100 m2) were compared with those obtained from two subsets with plots of 1–5 m2 and 6–30 m2. Results Most of the specialists were distributed among all the biogeographic regions, but many were more represented in the Alpine than in the Atlantic, Boreal and Continental regions. Variation in species composition was mainly correlated to temperature, precipitation and latitude in the three data sets, showing a major gradient from (1) alpine belt fens characterized by spring species to (2) small sedge fens mainly distributed in mountain regions and (3) boreo-temperate fens reflecting waterlogged conditions. Conclusions Base-rich fen communities are widely distributed across European biogeographic regions, but the Alpine region can be considered as the compositional centre of this vegetation type. Large-scale gradients of species composition are mainly explained by climate, while the influence of latitude is probably correlated to increasing water table in the boreo-temperate regions. These gradients can be better understood by differentiating three major vegetation types, which should be considered when establishing classification systems of base-rich fens in Europe.
Data management for the Braun-Blanquet project and the European vegetation archive
Jimenez-Alfaro, B. ; Hennekens, S.M. ; Chytry, M. - \ 2014
In: Proceedings of the Alaska Arctic Vegetation Archive Workshop. - Akureyri, Iceland : CAFF International Secretariat - ISBN 9789935431295 - p. 44 - 47.
The Braun-Blanquet project and the European Vegetation Archive are among the first initiatives for analyzing comprehensive datasets of vegetation plots in Europe (Jimenez-Alfaro et al. 2013). Both initiatives are based on the compilation of vegetation data from different collaborators, including national and regional databases and additional data from individual researchers or research groups or the literature. The management of this information is complex since it derives from heterogeneous sources and many different research contexts.
Review of EUNIS forest habitat classification
Schaminee, J.H.J. ; Chytry, M. ; Hennekens, S.M. ; Jiménez-Alfaro, B. ; Mucina, L. ; Rodwell, J.S. ; Tichy, R. - \ 2013
Copenhagen : EEA (EEA Technical Report EEA/NSV/13/005) - 111 p.
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