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The CLUE-S model is a popular choice for modelling land use and land cover change from local to regional scales, but it spatially allocates the demand for only the total cover of each land class in the predicted map. In the present work, we introduce a CLUE-S variant that allocates demand at the more detailed level of land type transitions, the trans-CLUE-S model. We implemented this extension algorithmically in R, without the need of new parameters. By processing each row of the land transition matrix separately, the model allocates the demand of each land category’s transitions via the CLUE-S allocation routine for only the cells which were of that category in the map of the previous time step. We found that the trans-CLUE-S model had half the total and configuration disagreement of the CLUE-S predictions in an empirical landscape, and in simulated landscapes of different characteristics. Moreover, the trans-CLUE-S performance was less sensitive to the number of environmental predictors of land type suitability for allocating demand. Although trans-CLUE-S is computationally more demanding due to running a CLUE-S allocation for each land class, we appended the solution of a land-use assignment optimisation problem that facilitates the convergence and acceleration of allocation. We additionally provide R functions for: CLUE-S variants at other levels of demand resolution; random instead of environment-based allocation; and for simulating landscapes of desired characteristics. Our R code for the models and functions can contribute to more reproducible, transparent and accurate modelling, analysis and interpretation of land cover change.HighlightsThe trans-CLUE-S model employs demand at the finer level of land type transitionsThe trans-CLUE-S predictions were twice more accurate than the CLUE-S model’sThe trans-CLUE-S accuracy was less dependent on the amount of environmental dataAlgorithmic addition of a land assignment task enabled and sped up full convergenceR code is provided for our models and auxiliary functions

Caio Graco-Roza was funded by the Coordination for the Improvement of Higher Education Personnel (CAPES), the Carlos Chagas Filho Research Support Foundation (FAPERJ) and the Ella and Georg Erhnrooth Foundation; Jan Altman by research grants INTER-EXCELLENCE LTAUSA19137 provided by Czech Ministry of Education, Youth and Sports, 20-05840Y of the Czech Science Foundation, and long-term research development project no. RVO 67985939 of the Czech Academy of Sciences; Otso Ovaskainen was funded by Academy of Finland (grant no. 309581), Jane and Aatos Erkko Foundation, Research Council of Norway through its Centres of Excellence Funding Scheme (223257), and the European Research Council (ERC) under the European Union's Horizon 2020 research and innovation programme (grant agreement no. 856506; ERC-synergy project LIFEPLAN); and Jianjun Wang was funded by CAS Key Research Program of Frontier Sciences (QYZDB-SSW-DQC043) and National Natural Science Foundation of China (91851117). The "sPlot" project was initiated by sDiv, the Synthesis Centre of the German Centre for Integrative Biodiversity Research (iDiv) Halle-Jena-Leipzig, funded by the German Research Foundation (DFG FZT 118), and is now a platform of iDiv. The study was supported by the TRY initiative on plant traits (). We are also grateful to Jens Kattge and TRY database. TRY is hosted, developed and maintained at the Max Planck Institute for Biogeochemistry (MPI-BGC) in Jena, Germany, in collaboration with the German Centre for Integrative Biodiversity Research (iDiv) Halle-Jena-Leipzig. The CESTES database of metacommunities is also an initiative of iDiv led by Alienor Jeliazkov. We thank sDiv for supporting the open science initiative., Aim: Understanding the variation in community composition and species abundances (i.e., β-diversity) is at the heart of community ecology. A common approach to examine β-diversity is to evaluate directional variation in community composition by measuring the decay in the similarity among pairs of communities along spatial or environmental distance. We provide the first global synthesis of taxonomic and functional distance decay along spatial and environmental distance by analysing 148 datasets comprising different types of organisms and environments. Location: Global. Time period: 1990 to present. Major taxa studied: From diatoms to mammals. Method: We measured the strength of the decay using ranked Mantel tests (Mantel r) and the rate of distance decay as the slope of an exponential fit using generalized linear models. We used null models to test whether functional similarity decays faster or slower than expected given the taxonomic decay along the spatial and environmental distance. We also unveiled the factors driving the rate of decay across the datasets, including latitude, spatial extent, realm and organismal features. Results: Taxonomic distance decay was stronger than functional distance decay along both spatial and environmental distance. Functional distance decay was random given the taxonomic distance decay. The rate of taxonomic and functional spatial distance decay was fastest in the datasets from mid-latitudes. Overall, datasets covering larger spatial extents showed a lower rate of decay along spatial distance but a higher rate of decay along environmental distance. Marine ecosystems had the slowest rate of decay along environmental distances. Main conclusions: In general, taxonomic distance decay is a useful tool for biogeographical research because it reflects dispersal-related factors in addition to species responses to climatic and environmental variables. Moreover, functional distance decay might be a cost-effective option for investigating community changes in heterogeneous environments., Coordenacao de Aperfeicoamento de Pessoal de Nivel Superior (CAPES) Fundacao Carlos Chagas Filho de Amparo a Pesquisa do Estado do Rio De Janeiro (FAPERJ), Ella and Georg Erhnrooth Foundation, Ministry of Education, Youth & Sports - Czech Republic LTAUSA19137 Grant Agency of the Czech Republic 20-05840Y Czech Academy of Sciences RVO 67985939, Academy of Finland 309581, Jane and Aatos Erkko Foundation, Research Council of Norway through its Centres of Excellence Funding Scheme 223257, European Research Council (ERC) 856506, CAS Key Research Program of Frontier Sciences QYZDB-SSW-DQC043, National Natural Science Foundation of China (NSFC) 91851117, German Research Foundation (DFG) DFG FZT 118, TRY initiative on plant traits

The size and shape of tree leaves and their variation within the canopy are the result of both physiological plasticity and an overall adaptive strategy against unfavourable environmental conditions. In this study, diversity patterns at leaf morphological traits will be described within and among populations of trees with different phylogenetic background. Beech (Fagus sp.) is a widespread tree in Eurasia, represented by two species; F. sylvatica in Europe and F. orientalis in eastern Europe and Asia. Both species appear in the Rodopi mountains, in southeast Balkans. Five beech populations were sampled in the southern slopes of Rodopi along a west–east gradient representing an established transitional zone between the two beech species. The diversity of six leaf traits was examined in shade leaves and leaves exposed to direct irradiation. Significant differences appeared among populations and among the two shading classes. Western beech populations consisted of trees with smaller leaves and fewer veins and were morphologically closer to F. sylvatica, while eastern populations seemed to be closer to F. orientalis. Shade leaves were constantly larger and less round than light leaves, probably due to different light harvesting strategies. The differences between populations were larger for shade leaves than for light leaves and presented a clear east–west trend, consistent to the differentiation pattern provided by previous genetic studies in the same region. Our results indicate that shade leaves probably maintain their size and shape independent from light irradiation and therefore may better express genetic differences among populations.

Studies comparing species’ distribution range at different time periods provide valuable information about temporal trends, and may help to identify the factors responsible for these changes. This knowledge is vital for setting conservation priorities and applying suitable management strategies and practices focused on species’ conservation. We examined records produced before and after December 31, 1985 for determining possible changes in the distribution range of 195 orchid species recorded in Greece, using a 10 × 10 km grid as the recording unit. We used regression techniques to explore the association of species richness with environmental and spatial variables. Species richness was calculated by using the orchid records made (a) before December 31, 1985, (b) after January 1, 1986, and (c) in both time periods. Generalized linear models (GLM) were used for identifying the most significant factors associated with the distribution of the threatened orchids of Greece. Most Greek orchids showed an increasing trend in distribution from the first to the second period, with the exception of a few Critically Endangered species, whose distribution was larger in the first than in the second period. The spatial distribution of the threatened orchids was mostly explained by land area and the area of the calcareous substrates at each grid cell, whereas all variables that were used in the GLM analysis accounted for about 32% of the variance. The observed increasing trend in orchid distribution should be attributed to the extensive research efforts during the second period, especially in areas of high altitude and towards the northern parts of Greece. Our results did not support the findings of other studies, mainly from northern Europe, which showed a significant decline in the distribution range of their orchid flora. These differences in distribution trends could be attributed to the complex orographic configuration of Greece, which renders large parts of the country rather unsuitable for agricultural use or for extensive urban development.

Hybridization is a process of great importance in the ecology and evolution of plants. However, identifying hybrids is not an easy task, especially in cases of sympatry. Such cases are the Abies taxa in Greece; Silver fir (A. alba), Greek fir (A. cephalonica) and King Boris fir (A. × borisii-regis). The latter has been characterized as a hybrid between the first two species as its morphological traits are either a combination of the parental ones or intermediate. Besides the morphological traits, A. × borisii-regis has so far been identified only at the population level, with biochemical and DNA markers. Here, we describe the identification of A. × borisii-regis at the individual level by employing a species-specific marker in the cpDNA and another in mtDNA, taking advantage of their uniparental inheritance mode from the paternal and the maternal parent, respectively. In addition, based on these markers, essential information emerges on the distribution and the mtDNA diversity of the Abies taxa in Greece.

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.

The EU Water Framework Directive foresees the ecological assessment of surface waters against identified pressures. Nutrient loading is the main pressure impairing the ecological quality of lake ecosystems, and aquatic macrophytes are considered good indicators of ecological response. In this study, we statistically assessed different aspects of aquatic plant (macrophyte) diversity in response to different trophic levels in Mediterranean lakes. We used 5690 relevés of aquatic vegetation, distributed over 305 transects, sampled in 18 freshwater lake ecosystems during 2013–2016. Our results show a significant decrease in taxonomic alpha diversity in lakes with a total phosphorus content above 100 μg/L. Syntaxonomic diversity followed the species richness pattern as well. Functional richness decreased along the trophic gradient, while functional dispersion was higher in lakes with high trophic levels. Taxonomic and functional beta partitioning presented changes in assembly processes leading to greater community homogeneity in lakes with higher trophic levels. In summary, we found no redundancy between taxonomic and functional diversity indices. These results provide novel insights into aquatic plant assembly processes of impacted freshwater lakes needed to forward conservation and restoration practices.

The neglected and underutilized plants (NUPs) could become alternative food sources in the agro-alimentary sector, enriching human and animal diets, offering the opportunity for sustainable exploitation, resilience to climate change, and production with resistance to pests and diseases. In the Mediterranean countries, these valuable resources are threatened by climate change, overexploitation, and/or monoculture. In this framework, we evaluated 399 local endemic NUPs of Crete (Greece), the Mediterranean coast, Rif of Morocco, and Tunisia, regarding their agro-alimentary potential, and assessed their feasibility and readiness timescale for sustainable exploitation with own previously published methodology. The methodological scheme was developed by experts in co-creative workshops, using point-scoring of seven attributes to evaluate the potential of the targeted NUPs in the agro-alimentary. Our results showed a diversity of promising local endemic NUPs of different families in the studied regions (Lamiaceae members are prominent), and we outlined the cases of 13 taxa with the highest optimum scores of agro-alimentary potential (&gt, 70%). Despite the diversity or the promising potential and current ex-situ conservation efforts to bridge gaps, our study indicated that only a few cases of Cretan local endemic NUPs can be sustainably exploited in the short-term. However, it is argued that many more local endemic NUPs can easily follow sustainable exploitation schemes if specific research gaps are bridged. Since NUPs can help to increased diversification of food production systems by adding new nutritional/beneficial species to human and animal diets, basic and applied research, as well as market and stakeholder attraction, is suggested as prerequisite to unlock the full potential of the focal endemic NUPs in the agro-alimentary sector.

Patterns of phylogenetic diversity and structure of flora in refugial areas may be very complex due to habitat heterogeneity and legacies of various historical processes. Revealing these patterns can help understand the nature of the refugia. Here we study phylogenetic patterns of the flora in one of the main putative Pleistocene refugia of temperate forest biota in Europe through a community-level analysis for the main types of forests. For this purpose, a database of vegetation plots from deciduous and mountain coniferous forests of northern and central Greece was created. The plots were classified into floristically and ecologically interpretable community types. Faith’s phylogenetic diversity, mean phylogenetic distance and mean nearest taxon distance, as well as their standardized equivalents, were used to characterize the mean phylogenetic diversity and structure of each community type. All analyses were applied both for all the vascular plant taxa and for angiosperm taxa only. We related the measures of phylogenetic diversity and structure to ecological factors and compared them among community types. The 25 identified community types differed in their phylogenetic diversity and structure. The observed patterns were significantly different when old evolutionary lineages, such as gymnosperms and ferns, were taken into account. A random or overdispersed phylogenetic structure was observed for most of the community types, while only two oak-dominated community types were clustered. The highest phylogenetic diversity and a notably high proportion of plots with an overdispersed phylogenetic structure were observed in two mesic ravine forest community types (one of these transitional to beech forests). Phylogenetic metrics correlated with ecological conditions, in particular, overdispersion was more common in shaded, cool and moist forests. The high phylogenetic diversity and overdispersed structure of the two ravine community types can be related to the putative refugial role of these forest habitats.

Neottia cordata is an orchid species that exclusively occurs in Picea abies subsp. abies and Pinus sylvestris stands and forms its southernmost distribution limits in Greece. Over the last decade, it has been recorded in numerous sites, in which it forms small colonies, both in terms of population and area. This research re-evaluates the threat status of Neottia cordata based on updated chorological and population data; it also checks the effectiveness of the Natura 2000 Network as far as its conservation is concerned by applying the MaxEnt model. The recently collected data justify the classification of N. cordata as "Vulnerable". The most significant variables that affect its distribution were found to be the vegetation type and the precipitation of the warmest quarter. The results of the MaxEnt model suggest that: (i) the Natura 2000 Network is characterised by areas of higher habitat suitability values when compared to the areas that fall outside this network; and, (ii) Picea abies subsp. abies forests are more suitable for its conservation than those of Pinus sylvestris. It is concluded that the most appropriate management measure for the species conservation is the maintenance of the tree layer canopy closed.

Questions: The human-related spread of alien plants has serious environmental and socioeconomic impacts. Therefore, it is important to know which habitats are most threatened by invasion and why. We studied a wide range of European grasslands to assess: (a) which alien species are the most successful invaders in grasslands; (b) how invasion levels differ across European regions (countries or their parts) and biogeographical regions; and (c) which habitat types are the most invaded. Location: Europe. Methods: We selected 97,411 grassland vegetation plots from the European Vegetation Archive (EVA) and assigned a native or alien status to each of the 8,212 vascular plant species found in these plots. We considered only neophytes (alien species introduced after 1500 AD), which we further divided according to their origin. We compared the levels of invasion using relative neophyte richness in the species pool, relative neophyte richness and cover per plot, and percentages of invaded plots among regions and habitats. Results: Only 536 species, representing 6.5% of all grassland vascular plant species, were classified as neophytes. These were mostly therophytes or hemicryptophytes with low habitat specificity. Most of them were present in very few plots, while only three species were recorded in more than 1% of all plots (Onobrychis viciifolia, Erigeron annuus and Erigeron canadensis). Although invasion levels were generally low, we found more invaded plots in the Boreal and Continental regions. When considering only non-European neophytes, the Pannonian region was the most invaded. Among different grassland habitats, sandy grasslands were most invaded, and alpine and oromediterranean grasslands least invaded. Conclusions: In general, natural and semi-natural European grasslands have relatively low levels of neophyte invasions compared with human-made habitats or alluvial forests, as well as with grasslands on other continents. The most typical neophytes invading European grasslands are species with broad ecological niches., IA, VK, JDa, MV, IK and MC were supported by the Czech Science Foundation (EXPRO grant no. 19-28491X), PP and JP by the Czech Science Foundation (EXPRO grant no. 19-28807X), JDa, PP and JP by the Czech Academy of Sciences (long-term research development project RVO 67985939), IB by the Basque Government (IT936-16), RGG by REMEDINAL network (S2018/EMT-4338), AL by Komarov Botanical Institute, RAS (AAAA-A19-119031290052-1) and AK by the National Research Foundation of Ukraine (grant no. 2020.01/0140). JCS considers this work a contribution to his VILLUM Investigator project “Biodiversity Dynamics in a Changing World” funded by VILLUM FONDEN (grant 16549).

Understanding the variation in community composition and species abundances, i.e., β-diversity, is at the heart of community ecology. A common approach to examine β-diversity is to evaluate directional turnover in community composition by measuring the decay in the similarity among pairs of communities along spatial or environmental distances. We provide the first global synthesis of taxonomic and functional distance decay along spatial and environmental distance by analysing 149 datasets comprising different types of organisms and environments. We modelled an exponential distance decay for each dataset using generalized linear models and extracted r2 and slope to analyse the strength and the rate of the decay. We studied whether taxonomic or functional similarity has stronger decay across the spatial and environmental distances. We also unveiled the factors driving the rate of decay across the datasets, including latitude, spatial extent, realm, and organismal features. Taxonomic distance decay was stronger along spatial and environmental distances compared with functional distance decay. The rate of taxonomic spatial distance decay was the fastest in the datasets from mid-latitudes while the rate of functional decay increased with latitude. Overall, datasets covering larger spatial extents showed a lower rate of decay along spatial distances but a higher rate of decay along environmental distances. Marine ecosystems had the slowest rate of decay. This synthesis is an important step towards a more holistic understanding of patterns and drivers of taxonomic and functional β-diversity.

The neglected and underutilized plants (NUPs) have never been mainstreamed by researchers, politicians and stakeholders for sustainable exploitation in the ornamental-horticultural sector. This study focused on 399 local endemic plants of three Mediterranean regions (Crete, Mediterranean coast-Rif of Morocco, Tunisia), with the aim to develop a new scheme for their multifaceted ornamental-horticultural evaluation facilitating their sustainable exploitation. The methodological scheme was developed within three multidisciplinary co-creative workshops by experts and was adjusted by end-users. The developed scheme uses point and weighted scoring of several attributes relevant to biological and ornamental-horticultural characteristics and concerns three ranking levels: (i) ornamental-horticultural potential (general or subsector-specific, 20 attributes), (ii) sustainable exploitation feasibility (12 attributes) and (iii) readiness timescale (after gap and SWOT analyses) in creating value chains in short-, medium- or long-term. The analysis of the data illustrated two example-cases of already achieved sustainable exploitation with established value chains, outlined the prospects for sustainable exploitation of 18 and 23 local endemic NUPs in short-term and medium-term, respectively, and identified 86 taxa with reachable possibilities in the long-term. The proposed multifaceted evaluation scheme can be applied for the valorisation of NUPs in other areas and may help to define priorities and to identify opportunities and gaps for their sustainable exploitation.

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.

Genista lydia Boiss. is an endemic Balkan-Anatolian species which forms rare communities in the territory of south Bulgaria and northeast Greece. They are spread exclusively on acidic, siliceous substrates. This study presents new data on their distribution, floristic and ecological structure and phytosociological affinities. The research is based on 156 phytosociological relevés. Unweighted pair–group method with arithmetic averages (UPGMA) was employed and a detrended correspondence analysis (DCA) was performed prior to the syntaxonomical decision. Three new associations and the new alliance Genistion lydiae have been described and classified within the order Lavandulo stoechadis-Hypericetalia olympici Mucina in Mucina et al. 2016 of the class Cisto-Lavanduletea stoechadis Br.-Bl. in Br.-Bl. et al. 1940. The study has also emphazised the potential threats concerning a decline of the habitat area and proposed some conservation measures.

Monitoring of habitat types conservation status is an essential task in the frame of the European policy for biodiversity conservation. The parameters to be assessed for the purposes of habitat types’ conservation status assessment are described in several European documents, but the methodology for their determination has not yet been standardized or optimized. This study presents methods for the assessment of the actual status and the future prospects of structure and functions of habitat types. Specifically, it presents a bottom–up approach for the assessment of these two parameters at different spatial scales. In the proposed method, conservation status assessment is based on a classification of habitat types to subtypes, with the latter representing the basic monitoring entities. The conservation status is assessed by recording: (i) the presence/absence of specific indicators of structure and functions per habitat type, and (ii) the presence/absence, abundance, and vitality of the typical species of the habitat subtypes. The typical species are determined objectively using algorithms and fidelity coefficient values. The conservation status and future prospects of structure and functions (including the typical species) are estimated quantitatively with the help of numerical methods and algorithms, but their assignment to conservation status classes is based on thresholds defined by experts. Assessments are made at the local scale, but can be upscaled to coarser ones (up to the national level). The proposed methods have been applied in Greece and were effective both in terms of results obtained and costs needed.

The goal of this study was to quantitatively assess the relationship linking vegetation and airborne pollen. For this, we established six sampling stations in the city of Thessaloniki, Greece. Once every week for 2 years, we recorded airborne pollen in them, at breast height, by use of a portable volumetric sampler. We also made a detailed analysis of the vegetation in each station by counting all existing individuals of the woody species contributing pollen to the air, in five zones of increasing size, from 4 to 40 ha. We found the local vegetation to be the driver of the spatial variation of pollen in the air of the city. Even at very neighbouring stations, only 500 m apart, considerable differences in vegetation composition were expressed in the pollen spectrum. We modelled the pollen concentration of each pollen taxon as a function of the abundance of the woody species corresponding to that taxon by use of a Generalized Linear Model. The relationship was significant for the five most abundantly represented taxa in the pollen spectrum of the city. It is estimated that every additional individual of Cupressaceae, Pinaceae, Platanus, Ulmus and Olea increases pollen in the air by approximately 0.7, 0.2, 2, 6 and 5%, respectively. Whether the relationships detected for the above pollen taxa hold outside the domain for which we have data, as well as under different environmental conditions and/or with different assemblages of species representing them are issues to be explored in the future.

We studied the differentiation among plant communities of deciduous broadleaved and mountain coniferous forests in terms of functional diversity and identity at a regional scale (northern and central Greece). We asked if patterns of functional differentiation among communities are consistent between the overstorey and understorey layers and if they can be influenced by deep past environmental conditions. Functional Richness (FRic) and Functional Dispersion (FDis), as well as their standardized effect sizes, were employed to assess the multivariate functional diversity of the community types. In contrast, single-trait Community Weighted Means (CWMs) were used as surrogates of functional identity. The aforementioned indices were calculated for three datasets, namely all the vascular plant taxa found in individual vegetation plots (total community), all phanerophyte (tree and shrub) taxa (overstorey) and all non-phanerophyte vascular plant taxa (understorey). We found that community types and especially four broad forest types (beech, ravine, pine and oak forests) are well differentiated in terms of functional composition (identity), as indicated by Non-Metric Multidimensional Scaling (NMDS). After conducting an NMDS for the three datasets, functional identity based on the total floristic composition was found to be the best discriminator of the studied communities. However, contrasting patterns were found for some specific traits or their categories between overstorey and understorey layers. The patterns of functional diversity of the community types (based on multivariate indices), revealed by calculating the standardized effect sizes of FRic and FDis based on the richness null model, did not differ substantially from random expectations for most of the studied community types when the dataset of all the vascular plant taxa was analyzed. However, the patterns revealed for the overstorey layer differed from those for the understorey layer. For the latter layer, the clustered structure was revealed in many community types based on the ses.FDis metric. Indications of deep past influence on the functional composition were found for certain community types (i.e. ravine forests) based on single-trait metrics, but no indication of such influence was found based on multivariate indices. Our findings highlight the complementarity and the additive explanatory value of the simultaneous use of single- and multi-trait approaches and their application to different layers in forests.

Flowering and pollen seasons are sensitive to environmental variability and are considered climate change indicators. However, it has not been concluded to what extent flowering phenology is indeed reflected in airborne pollen season locally. The aim of this study was to investigate, for a wide spectrum of plant species, the responsiveness of flowering to different environmental regimes and also to check for commensurate changes in the respective pollen seasons. Fourteen taxa were selected representing the families of Corylaceae, Cupressaceae, Fagaceae, Oleaceae, Pinaceae and Platanaceae. Each was studied for at least one full flowering year and in at least two sampling stations differing in elevation or direction within the period 2004–2011. Flowering observations were made 2–3 times a week, from flower differentiation to flower desiccation; >1000 daily observations were made in 225 plant individuals, in total. Data were regressed against meteorological parameters in an attempt to identify the driving factors of flowering onset and offset. Flowering dates were compared with dates of the related airborne pollen seasons per taxon; airborne pollen monitoring took place daily using a Hirst-type volumetric sampler. For all taxa, flowering started earlier at lower elevations or southern direction. Cumulative temperature proved to be the factor most significantly affecting flowering onset and end for all species (p 0.70). For the majority of the taxa examined, airborne pollen season did not coincide locally with the respective flowering periods of the selected species: flowering could precede pollen season, even for more than one month (as in C. avellana), or start after the pollen season onset (as in P. brutia). The strong sensitivity of flowering to air temperature is not the case in the respective pollen seasons. This raises questions on the relationship between flowering times and airborne pollen seasons and on the rather underestimated role of long-distance transport of pollen.

This study presents a first, national scale approach on ecosystem condition assessment for Greece, through integrating the available surveillance and monitoring data for habitat types, at the plot level, within the Natura 2000 network. The study consists of two parts: (a) ecosystem condition assessment for ecosystem types in Greece, using the conservation degree at plot level as an indicator and (b) a large scale analysis of the forest types' condition using, as indicators, the pressures and typical plant Typical species richness, as an exemplary case to interpret the outcomes of the assessment. The main results of this study revealed that: (i) the majority of the ecosystem types are in above good condition, with the higher percentages of bad condition recorded for wetlands, rivers and lakes, marine inlets and transitional waters, (ii) forest categories in their majority are at above-adequate condition, (iii) at forest ecosystem categories specific pressures (e.g. grazing, cultivations, forestry clearance) act as main drivers forecosystem condition change, (iv) Mediterranean deciduous forests are the most floristic-rich forest category, regarding typical plant Typical species. Simultaneously, it is highlighted that already available datasets could be used for immediate and rapid framework assessments, which will guide future steps on ES studies, research and decision-making.

The Water Framework Directive (WFD) requires Member States to develop appropriate assessment methods for the classification of the ecological status of their surface waters. Mediterranean region has lagged behind in this task, so we propose here the first developed method for Greek lakes, Hellenic Lake Macrophyte (HeLM) assessment method. This method is based on two metrics, a modified trophic index and maximum colonization depth Cmax that quantify the degree of changes in lake macrophytic vegetation, as a response to eutrophication and general degradation pressures. The method was developed on the basis of a data set sampled from 272 monitoring transects in 16 Greek lakes. Sites from three lakes were selected as potential reference sites by using a screening process. Ecological quality ratios were calculated for each metric and for each lake, and ecological status class boundaries were defined. For the evaluation of effectiveness of the method, the correlations between individual metrics and final HeLM values and common pressure indicators, such as total phosphorus, chlorophyll a and Secchi depth, were tested and found highly significant and relatively strong. In addition, the ability of HeLM values and its individual metrics to distinguish between different macrophytic communities’ structure was checked using aquatic plant life-forms and found satisfactory. The HeLM method gave a reliable assessment of the macrophytic vegetation’s condition in Greek lakes and may constitute a useful tool for the classification of ecological status of other Mediterranean lakes. Electronic supplementary material The online version of this article (10.1007/s10661-018-6708-1) contains supplementary material, which is available to authorized users.

International audience; A central hypothesis of ecology states that regional diversity influences local diversity through species-pool effects. Species pools are supposedly shaped by large-scale factors and then filtered into ecological communities, but understanding these processes requires the analysis of large datasets across several regions. Here, we use a framework of community assembly at a continental scale to test the relative influence of historical and environmental drivers, in combination with regional or local species pools, on community species richness and community completeness. Using 42,173 vegetation plots sampled across European beech forests, we found that large-scale factors largely accounted for species pool sizes. At the regional scale, main predictors reflected historical contingencies related to post-glacial dispersal routes, whereas at the local scale, the influence of environmental filters was predominant. Proximity to Quaternary refugia and high precipitation were the main factors supporting community species richness, especially among beech forest specialist plants. Models for community completeness indicate the influence of large-scale factors, further suggesting community saturation as a result of dispersal limitation or biotic interactions. Our results empirically demonstrate how historical factors complement environmental gradients to provide a better understanding of biodiversity patterns across multiple regions.

International audience; 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.

International audience; Plant functional traits directly affect ecosystem functions. At the species level, trait combinations depend on trade-offs representing different ecological strategies, but at the community level trait combinations are expected to be decoupled from these trade-offs because different strategies can facilitate co-existence within communities. A key question is to what extent community-level trait composition is globally filtered and how well it is related to global versus local environmental drivers. Here, we perform a global, plot-level analysis of trait-environment relationships, using a database with more than 1.1 million vegetation plots and 26,632 plant species with trait information. Although we found a strong filtering of 17 functional traits, similar climate and soil conditions support communities differing greatly in mean trait values. The two main community trait axes that capture half of the global trait variation (plant stature and resource acquisitiveness) reflect the trade-offs at the species level but are weakly associated with climate and soil conditions at the global scale. Similarly, within-plot trait variation does not vary systematically with macro-environment. Our results indicate that, at fine spatial grain, macro-environmental drivers are much less important for functional trait composition than has been assumed from floristic analyses restricted to co-occurrence in large grid cells. Instead, trait combinations seem to be predominantly filtered by local-scale factors such as disturbance, fine-scale soil conditions, niche partitioning and biotic interactions.

GrassPlot is a collaborative vegetation-plot database organised by the Eurasian Dry Grassland Group (EDGG) and listed in the Global Index of Vegetation-Plot Databases (GIVD ID EU-00-003). GrassPlot collects plot records (relevés) from grasslands and other open habitats of the Palaearctic biogeographic realm. It focuses on precisely delimited plots of eight standard grain sizes (0.0001; 0.001; ... 1,000 m²) and on nested-plot series with at least four different grain sizes. The usage of GrassPlot is regulated through Bylaws that intend to balance the interests of data contributors and data users. The current version (v. 1.00) contains data for approximately 170,000 plots of different sizes and 2,800 nested-plot series. The key components are richness data and metadata. However, most included datasets also encompass compositional data. About 14,000 plots have near-complete records of terricolous bryophytes and lichens in addition to vascular plants. At present, GrassPlot contains data from 36 countries throughout the Palaearctic, spread across elevational gradients and major grassland types. GrassPlot with its multi-scale and multi-taxon focus complements the larger international vegetation-plot databases, such as the European Vegetation Archive (EVA) and the global database “sPlot”. Its main aim is to facilitate studies on the scale- and taxon-dependency of biodiversity patterns and drivers along macroecological gradients. GrassPlot is a dynamic database and will expand through new data collection coordinated by the elected Governing Board. We invite researchers with suitable data to join GrassPlot. Researchers with project ideas addressable with GrassPlot data are welcome to submit proposals to the Governing Board. Abbreviations: EDGG = Eurasian Dry Grassland Group; EVA = European Vegetation Archive; GrassPlot = Database of Scale-Dependent Phytodiversity Patterns in Palaearctic Grasslands; SAR = species-area relationship.

Changes in species diversity following volcanic eruptions have been studied extensively, but our knowledge on functional diversity and community assembly under such conditions is very limited. Here, we study the processes following the destruction of vegetation after a volcanic eruption. Specifically, we investigate (1) the temporal patterns of taxonomic and functional diversity over time since a previous eruption (alpha diversity) and beta diversity, (2) the temporal patterns of 26 individual traits (vegetative characteristics, plant taxa ecological preferences, and regenerative characteristics) providing more detailed information on species strategies at the initial and later stages of succession, and (3) the processes driving species assembly and whether they changed over time since the eruption an eruption. We analyzed data recorded during five floristic censuses that took place between 1911 and 2011, calculated alpha and beta facets of taxonomic and functional diversity and examined how community structure changed over time, using 26 functional characteristics, based on their ability to discern primary from later colonists, including longevity, growth form, Ellenberg's indicator values, seed production and weight, flower size and sex, pollination type, and dispersal mode. Null model analysis was used to test whether the observed functional diversity deviates from random expectations. Alpha diversity, both taxonomic and functional, increased over time after an eruption, while beta diversity did not display a clear trend. This finding indicates that mainly abiotic processes determine species assembly over time after an eruption (at least for the time span studied here), contrary to theoretical expectations. It is most interesting that, simultaneously, some aspects of diversity indicated the effect of biotic interactions (facilitation and competition) on the assembly of species a few years after an eruption. This finding implies a legacy effect, since a high percentage of perennial species was noticed in the assemblage right after the eruption, as well as the effect of the harsh environmental conditions on the assembly of the plant communities. In conclusion, our results indicate the role of legacy effects in succession (most probably through the survival of underground plant parts) and underline the importance of disturbance history in providing the context needed for understanding effects of past events on succession.

Aim: Formalized classifications synthesizing vegetation data at the continental scale are being attempted only now, although they are of key importance for nature conservation planning. Therefore, we aim to provide a vegetation classification and to describe the main biogeographical patterns of floodplain forests and alder carrs in Europe. Location: Europe. Methods: A database of more than 40 000 vegetation plots of floodplain forests and alder carrs across Europe was compiled. After geographic stratification, 16 392 plots were available for classification, which was performed using the supervised method Cocktail. We also searched for new associations using semi-supervised K-means classification. The main biogeographic patterns and climate-related gradients in species composition were determined using detrended correspondence analysis and cluster analysis. Results: Thirty associations of floodplain forests and alder carrs were distinguished, which belong to five alliances. The Alnion incanae includes riparian, seepage and hardwood floodplain forests in the nemoral and hemiboreal zones (dominated by Alnus glutinosa and Fraxinus excelsior) and in the boreal zone (dominated by A. incana). The Osmundo-Alnion represents oceanic vegetation dominated by Alnus glutinosa, Fraxinus angustifolia and F. excelsior distributed mostly on the Iberian Peninsula and composed of species with Atlantic distribution and Iberian endemics. The Populion albae comprises floodplain forests frequently dominated by Fraxinus angustifolia, Populus alba and P. nigra that are widespread in floodplains of large rivers under summer-dry climates in the Mediterranean region. The Platanion orientalis represents eastern Mediterranean floodplain forests dominated by Platanus orientalis. The Alnion glutinosae includes forest swamps dominated by Alnus glutinosa distributed mostly in the nemoral and hemiboreal zones. The main biogeographic patterns within European floodplain forests and alder carrs reflect the climatic contrasts between the Mediterranean, nemoral, boreal and mountain regions. Oceanic floodplain forests differ from those in the rest of Europe. The hydrological regime appears to be the most important factor influencing species composition within regions. Conclusions: This study is the first applying a formalized classification at the association level for a broad vegetation type at the continental scale. The proposed classification provides the scientific basis for the necessary improvement of the habitat classification systems used in European nature conservation.

Premise of research. Even though the Balkan Peninsula is a biodiversity hot spot in southeastern Europe harboring many endemic plants, very little is known about the temporal extent of the phylogeographic history and the contemporary genetic diversity of the endemics there. Haberlea rhodopensis is one of the European Gesneriaceae species occurring in this region and represents appropriate study material to address these questions.Methodology. We generated fossil-dated molecular phylogenies (atpB-rbcL, trnH-psbA, trnLF, ITS) across the Lamiales to determine the origin and age of H. rhodopensis and conducted phylogeographic (trnH-psbA, ITS) and population genetic (ISSRs) analyses on 17 populations from Bulgaria and Greece, covering the entire distribution range of the species, to investigate their biogeographic history, present-day genetic diversity, and differentiation levels.Pivotal results. The European Gesneriaceae genera have a Tertiary origin in the early Oligocene, while the Haberlea lineage emerged ...

Questions Do the mechanisms driving community assembly differ between two islands of different age and history of vegetation development? How does sampling scale affect the strength that each assembly mechanism poses on the assembly of plant communities? Location Volcanic islands (Palea Kameni and Nea Kameni) of Santorini Archipelago, Greece. Methods Functional diversity has been proposed as a framework for discriminating among mechanisms of community assembly, such as habitat filtering, limiting similarity and random assembly. We investigated four plant communities in two sea-born volcanic islands. We recorded plant diversity at scales from 1 m2 to 64 m2. We calculated three indices of functional diversity: functional richness, functional evenness and functional divergence, using 26 functional traits (including vegetative characteristics, ecological preferences and regenerative characteristics). We used null model analysis to test for two different assembly mechanisms: habitat filtering and limiting similarity or random assembly. Results The assemblage of the four communities was complex and did not follow a single mechanism. In most cases, finer-scale patterns indicated randomness, while coarser scales revealed more structured communities. In the older island, the scrub community was mainly defined by limiting similarity. The therophytic community displayed a limited range of functional traits, indicating mainly habitat filtering, but within this range, the evenness of the distribution indicated limiting similarity. On the younger island, the range of traits did not differ from random. However, within this range of traits, one therophytic community showed signs of limiting similarity, while the second therophytic community displayed uneven functional trait distribution, indicating mainly habitat filtering. The three indices reflected different facets of functional diversity and were not correlated, thus we may argue that they are not redundant, and we even detected different mechanisms of assembly within the same community. Conclusions The functional diversity of the therophytic communities in the younger island implied no specific assembly mechanism; perhaps due to its age, the community is still at the early stages of colonization (i.e. stochastic processes, such as arrival of new species, prevail). In the older island, the lack of disturbances for a long period allowed the establishment of communities assembled by specific mechanisms, such as competition and habitat filtering.

QuestionsWhat are the main floristic patterns in European beech forests? Which classification at the alliance and suballiance level is the most convincing?LocationEurope and Asia Minor.MethodsWe applied a TWINSPAN classification to a data set of 24605 releves 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.ResultsThe 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.ConclusionsWe 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.