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Aims: The majority of research on the diversity-productivity relationship has utilized aboveground plant biomass as a measure of overall ecosystem productivity and investigated successions with species-poor early stages. During this study we modelled patterns of plant diversity and carbon pools along a succession from pasture abandonment to submediterranean Quercus cerris woodland and investigated ecosystem productivity using biomass and soil carbon. Study area: The hilly-mountainous area of Tolfa in Central Italy. Methods: We used chronosequences to analyse changes in vascular plant species richness and composition across successional stages and along a gradient of productivity (i.e. of increasing carbon stocks). To test possible differences across successional stages species richness and carbon pool data were analysed through analysis of variance. Mixed model regression analyses were used to test the effect of the amount of the carbon stocks on the variation in plant species richness and composition. Results: We found a high variation in plant species richness along the succession, with grasslands and newly formed woods showing the highest values. The amount of carbon in biomass and soil increased along the succession, even if soil carbon did not differ significantly among successional stages. Plant species richness was not dependent on the variation in the carbon pools (i.e. biomass and soil carbon) along the succession, while plant species composition significantly responded to this variation. Conclusions: Our results strongly support the need to manage semi-natural ecosystems through a multi-functional perspective that values both carbon sequestration and biodiversity. [ABSTRACT FROM AUTHOR]

The European biodiversity and forest strategies rely on forest sustainable management (SFM) to conserve forest biodiversity. However, current sustainability assessments hardly account for direct biodiversity indicators. We focused on forest multi-taxon biodiversity to: i) gather and map the existing information; ii) identify knowledge and research gaps; iii) discuss its research potential. We established a research network to fit data on species, standing trees, lying deadwood and sampling unit description from 34 local datasets across 3591 sampling units. A total of 8724 species were represented, with the share of common and rare species varying across taxonomic classes: some included many species with several rare ones (e.g., Insecta); others (e.g., Bryopsida) were represented by few common species. Tree-related structural attributes were sampled in a subset of sampling units (2889; 2356; 2309 and 1388 respectively for diameter, height, deadwood and microhabitats). Overall, multi-taxon studies are biased towards mature forests and may underrepresent the species related to other developmental phases. European forest compositional categories were all represented, but beech forests were over-represented as compared to thermophilous and boreal forests. Most sampling units (94%) were referred to a habitat type of conservation concern. Existing information may support European conservation and SFM strategies in: (i) methodological harmonization and coordinated monitoring; (ii) definition and testing of SFM indicators and thresholds; (iii) data-driven assessment of the effects of environmental and management drivers on multi-taxon forest biological and functional diversity, (iv) multi-scale forest monitoring integrating in-situ and remotely sensed information. • Through a Europe-wide research network we harmonized and merged 34 datasets on forest biodiversity • Multi-taxon biodiversity information is available for studied across 3,591 sampling units • Available data unevenly cover the European forest compositional categories • Knowledge gaps are boreal, Mediterranean, thermophilous and floodplain forests • The available information has the potential to inform conservation and management strategies for European forests [ABSTRACT FROM AUTHOR]

Aim: Primary forests have high conservation value but are rare in Europe due to historic land use. Yet many primary forest patches remain unmapped, and it is unclear to what extent they are effectively protected. Our aim was to (1) compile the most comprehensive European‐scale map of currently known primary forests, (2) analyse the spatial determinants characterizing their location and (3) locate areas where so far unmapped primary forests likely occur. Location: Europe. Methods: We aggregated data from a literature review, online questionnaires and 32 datasets of primary forests. We used boosted regression trees to explore which biophysical, socio‐economic and forest‐related variables explain the current distribution of primary forests. Finally, we predicted and mapped the relative likelihood of primary forest occurrence at a 1‐km resolution across Europe. Results: Data on primary forests were frequently incomplete or inconsistent among countries. Known primary forests covered 1.4 Mha in 32 countries (0.7% of Europe's forest area). Most of these forests were protected (89%), but only 46% of them strictly. Primary forests mostly occurred in mountain and boreal areas and were unevenly distributed across countries, biogeographical regions and forest types. Unmapped primary forests likely occur in the least accessible and populated areas, where forests cover a greater share of land, but wood demand historically has been low. Main conclusions: Despite their outstanding conservation value, primary forests are rare and their current distribution is the result of centuries of land use and forest management. The conservation outlook for primary forests is uncertain as many are not strictly protected and most are small and fragmented, making them prone to extinction debt and human disturbance. Predicting where unmapped primary forests likely occur could guide conservation efforts, especially in Eastern Europe where large areas of primary forest still exist but are being lost at an alarming pace. [ABSTRACT FROM AUTHOR]

Abstract: Questions: Do vascular plant species richness and beta‐diversity differ between managed and structurally complex unmanaged stands? To what extent do species richness and beta‐diversity relate to forest structural attributes and heterogeneity? Location: Five national parks in central and southern Italy. Methods: We sampled vascular plant species composition and forest structural attributes in eight unmanaged temperate mesic forest stands dominated or co‐dominated by beech, and in eight comparison stands managed as high forests with similar environmental features. We compared plant species richness, composition and beta‐diversity across pairs of stands (unmanaged vs managed) using GLMMs. Beta‐diversity was quantified both at the scale of each pair of stands using plot‐to‐plot dissimilarity matrices (species turnover), and across the whole data set, considering the distance in the multivariate species space of individual plots from their centroid within the same stand (compositional heterogeneity). We modelled the relationship between species diversity (richness and beta‐diversity) and forest structural heterogeneity and individual structural variables using GLMMs and multiple regression on distance matrices. Results: Species composition differed significantly between managed and unmanaged stands, but not richness and beta‐diversity. We found weak evidence that plant species richness increased with increasing levels of structural heterogeneity and canopy diversification. At the scale of individual stands, species turnover was explained by different variables in distinct stands, with variables related to deadwood quantity and quality being selected most often. We did not find support for the hypothesis that compositional heterogeneity varies as a function of forest structural characteristics at the scale of the whole data set. Conclusions: Structurally complex unmanaged stands have a distinct herb layer species composition from that of mature stands in similar environmental conditions. Nevertheless, we did not find significantly higher levels of vascular plant species richness and beta‐diversity in unmanaged stands. Beta‐diversity was related to patterns of deadwood accumulation, while for species richness the evidence that it increases with increasing levels of canopy diversification was weak. These results suggest that emulating natural disturbance, and favouring deadwood accumulation and canopy diversification may benefit some, but not all, facets of plant species diversity in Apennine beech forests. [ABSTRACT FROM AUTHOR]

Questions To what extent have semi-natural grassland plant species composition and functional traits changed during the past five decades? Which are the main directions of change? Which environmental, management and landscape factors have determined these changes? Location Central and southern Apennines of Italy. Methods In 2013-2014, we used a stratified random sampling design to revisit 132 semi-natural grasslands originally sampled between 1966 and 1992. The exact locations of the historical plots were not available but they were associated with detailed vegetation maps. Plots were divided in 17 groups based on study area and original community type. To assess changes in species composition we ran PERMANOVA and RDA. The extent of change was estimated by comparing each new plot with the quantitative species pool of the corresponding historical group; species composition changes were calculated using Bray-Curtis coefficient, and shifts in functional traits were quantified as differences in community-weighted means. These measures of change were used as response variables in mixed effects models including climate, topography, management and landscape metrics as explanatory variables. Results The largest changes were related to successional dynamics, which occurred mainly in grasslands developing at low altitude and in the flattest sites where the soil organic horizon was relatively thick. Colonization by shrubs and trees was facilitated by high soil pH and amount of adjacent forests. Also, management factors affected the patterns of shifts in species composition and functional traits: grazing intensity was negatively related to the degree of compositional change as well as to the abundance of woody species, whereas the increase of species associated with frequent grazing disturbance was found at sites with higher values of soil salinity. Conclusions Apennine semi-natural grasslands have undergone substantial changes in species composition and functional traits during recent decades. Compositional changes mainly indicated successional dynamics, likely following a decrease in grazing intensity, whereas some more accessible grasslands experienced an increase in disturbance-tolerant species. The combination of the re-visitation approach with functional traits allowed us to clarify different directions of changes in species composition, adding information on the ecological processes related to these changes. [ABSTRACT FROM AUTHOR]

The extension of forest area is a globally accepted tool to offset CO 2 emissions from deforestation and the combustion of fossil fuels. The common assumption is that in addition to the perceived climate benefits increasing forest area will also support biodiversity, thus making afforestation a “win-win scenario”. Based on the existing scientific evidences, we show that joined climate and biodiversity benefits are strongly context-dependent and that the outcome of afforestation is often highly questionable. In Europe, grasslands managed at low intensity contribute substantially to biodiversity conservation and carbon storage. However, many of these grasslands have been lost due to abandonment and subsequent spontaneous succession towards woody vegetation, or due to land use intensification. Moreover, grasslands are the ecosystems most often deliberately afforested in the context of EU carbon-centered policies that may thus counteract biodiversity conservation programmes. By reviewing the main EU policies targeting forests and grasslands, we found a striking ambivalence between policies and funding schemes addressing grassland conservation on the one hand (e.g. Habitats Directive, green payments within the Common Agricultural Policy) and those supporting afforestation on the other (e.g. rural development funds). We suggest three measures towards a better harmonization of the European Union policies that target forest and grassland ecosystems: (1) promoting the alignment of the decisions taken across different policy sectors; (2) focusing on the whole range of ecosystem services and biodiversity issues rather than on carbon management only; (3) valuing systems managed at low-intensity for their multifunctionality. [ABSTRACT FROM AUTHOR]

A broad interest exists in developing structure-based indicators to use as proxies for other attributes that are difficult to assess, such as biological diversity. Summary variables that account for stand-scale forest structural complexity could facilitate the comparison among stands and provide a means of ranking stands in terms of their potential contribution to biodiversity. We developed an index of structural heterogeneity (SHI) for beech forests in southern Italy: (i) we established a preliminary list of 23 structural variables obtained from data routinely collected in forest inventories; (ii) we quantified these variables in a set of 64 beech-dominated stands encompassing a wide range of variability in the Cilento, Vallo di Diano and Alburni National Park; (iii) we identified a core set of attributes that take into account the main sources of structural heterogeneity identified in reference old-growth forests; and (iv) we combined these core attributes into a simple additive index (SHI). We identified eight core attributes that were rescaled to the range 0 to 10 using regression equations based on raw attribute data. The SHI was calculated as the sum of these attribute scores and then expressed as a percentage. The index performance was evaluated against ten reference old-growth beech stands in the Apennines. The index ranged between 38 and 79.1 (median=59.4) and was distributed normally for the calibration dataset. The SHI successfully discriminated between old-growth (range=71.9-99.9, median=85.1) and early-mature to mature forests. Furthermore, the SHI linearly increased with stand age and was higher in multi-layer high forests than in single- and double-layer forests. However, a large variation was detected within both management types and age classes. SHI could be helpful for foresters as a tool for quantifying and comparing structural heterogeneity before and after a silvicultural intervention aimed at restoring the structural complexity in second-growth stands. [ABSTRACT FROM AUTHOR]

Different assembly processes may simultaneously affect local-scale variation of species composition in temperate old-growth forests. Ground layer species diversity reflects chance colonization and persistence of low-dispersal species, as well as fine-scale environmental heterogeneity. The latter depends on both purely abiotic factors, such as soil properties and topography, and factors primarily determined by overstorey structure, such as light availability. Understanding the degree to which plant diversity in old-growth forests is associated with structural heterogeneity and/or to dispersal limitation will help assessing the effectiveness of silvicultural practices that recreate old-growth patterns and structures for the conservation or restoration of plant diversity. We used a nested sampling design to assess fine-scale species turnover, i.e. the proportion of species composition that changes among sampling units, across 11 beech-dominated old-growth forests in Southern Europe. For each stand, we also measured a wide range of environmental and structural variables that might explain ground layer species turnover. Our aim was to quantify the relative importance of dispersal limitation in comparison to that of stand structural heterogeneity while controlling for other sources of environmental heterogeneity. For this purpose, we used multiple regression on distance matrices at the within-stand extent, and mixed effect models at the extent of the whole dataset. Species turnover was best predicted by structural and environmental heterogeneity, especially by differences in light availability and in topsoil nutrient concentration and texture. Spatial distances were significant only in four out of eleven stands with a relatively low explanatory power. This suggests that structural heterogeneity is a more important driver of local-scale ground layer species turnover than dispersal limitation in southern European old-growth beech forests. [ABSTRACT FROM AUTHOR]

Questions What are the main drivers of variation in beta-diversity for Bromus erectus semi-natural dry grasslands of habitat 6210(*) at different scales? How should environmental variables and spatial patterns be taken into account to conserve the maximum possible beta-diversity within the habitat? Location Central Italy. Methods We used 195 vegetation relevés distributed in three nested extents: a single mountain, a mountain chain and southern Lazio. Multiple regression on distance matrices was performed using dissimilarity matrices based on: (1) species abundances as response variables; (2) spatial coordinates and environmental parameters (altitude, slope, percentage of rock and stone coverage, aspect, annual rainfall) as explanatory variables. The two groups of explanatory variables were used separately to partition the variation, and jointly to assess the relative contribution of each individual variable. Those variables found to significantly affect beta-diversity were used to: (1) compare beta-diversity levels between a set of randomly selected and a set of stratified relevés; and (2) analyse the habitat distribution across environmental gradients. These analyses, together with the curves describing the relationships between spatial distances and composition dissimilarities, were used to inform management decisions for the habitat. Results Most of the variance was explained by environmental variables, whose share was higher in the smallest and intermediate extent than in the broadest extent. Community dissimilarity increased in proportion to differences in altitude and spatial distances at every extent. Accordingly, at all the extents, the selection of relevés stratified by altitude or selected taking into account a minimum spatial distance included significantly higher levels of within-habitat beta-diversity, than randomly selected relevés. The relation of beta-diversity to the variation in aspect and annual rainfall varied at different extents. Conclusions Our results demonstrate that dry grassland management plans aimed at conserving the maximum within-habitat beta-diversity should take into account variation in environmental variables, among which altitude proved to be a critical factor at every extent. Also, spatial distances positively affect within-habitat beta-diversity levels, and scale-dependent minimum distances among habitat patches should be taken into account when selecting patches of habitat 6210(*) to be conserved in the study area. [ABSTRACT FROM AUTHOR]

[Display omitted] • Forest multi-taxon studies have similar aims but different sampling protocols. • The most sampled taxonomic groups are plants, beetles, lichens, birds, fungi. • Soil, litter and canopy resulted as undersampled forest habitats. • Sampling units and substrates differed widely within and among taxonomic groups. • Sampling methods for stand structure were relatively homogeneous. Forests host most terrestrial biodiversity and their sustainable management is crucial to halt biodiversity loss. Although scientific evidence indicates that sustainable forest management (SFM) should be assessed by monitoring multi-taxon biodiversity, most current SFM criteria and indicators account only for trees or consider indirect biodiversity proxies. Several projects performed multi-taxon sampling to investigate the effects of forest management on biodiversity, but the large variability of their sampling approaches hampers the identification of general trends, and limits broad-scale inference for designing SFM. Here we address the need of common sampling protocols for forest structure and multi-taxon biodiversity to be used at broad spatial scales. We established a network of researchers involved in 41 projects on forest multi-taxon biodiversity across 13 European countries. The network data structure comprised the assessment of at least three taxa, and the measurement of forest stand structure in the same plots or stands. We mapped the sampling approaches to multi-taxon biodiversity, standing trees and deadwood, and used this overview to provide operational answers to two simple, yet crucial, questions: what to sample? How to sample? The most commonly sampled taxonomic groups are vascular plants (83% of datasets), beetles (80%), lichens (66%), birds (66%), fungi (61%), bryophytes (49%). They cover different forest structures and habitats, with a limited focus on soil, litter and forest canopy. Notwithstanding the common goal of assessing forest management effects on biodiversity, sampling approaches differed widely within and among taxonomic groups. Differences derive from sampling units (plots size, use of stand vs. plot scale), and from the focus on different substrates or functional groups of organisms. Sampling methods for standing trees and lying deadwood were relatively homogeneous and focused on volume calculations, but with a great variability in sampling units and diameter thresholds. We developed a handbook of sampling methods (SI 3) aimed at the greatest possible comparability across taxonomic groups and studies as a basis for European-wide biodiversity monitoring programs, robust understanding of biodiversity response to forest structure and management, and the identification of direct indicators of SFM. [ABSTRACT FROM AUTHOR]

Forest composition is a faithful indicator of the stressors and disturbances that influence forest ecosystems, and it should be accounted for in Sustainable Forest Management policies. Indeed, the classification of forest ecosystems in forest types is considered as a key tool to improve the assessment and monitoring of forest biological diversity, and for the definition of management guidelines. Accordingly, the Ministerial Conference on the Protection of Forests in Europe has recognized the need of developing a pan-European forest classification in forest types, and has identified indicators of Sustainable Forest Management that should be applied by forest types. The classification of vegetation has always been among the main aims of the plant sociology. The quantitative and qualitative analysis of plant species composition, performed through the plant sociological approach, condenses compositional and structural information within a hierarchical system, and expresses all historical, sociological and habitat factors that influence the actual and potential vegetation. In a modern perspective the integration of plant sociology and ecological analysis represents a key to a hierarchical land classification and to the understanding of vegetation dynamics; furthermore the long history of plant sociology determined the availability of large datasets of vegetation data throughout Europe. Starting from these considerations, in this paper we briefly describe how plant sociology could represent a tool for the assessment of the indicators of SFM that should be applied by forest types, giving insights on how this discipline could contribute to the assessment of each of these indicators. [ABSTRACT FROM AUTHOR]

Question: Indices of functional diversity have been seen as the key for integrating information on species richness with measures that focus on those components of community composition related to ecosystem functioning. For comparing species richness among habitats on an equal-effort basis, so-called sample-based rarefaction curves may be used. Given a study area that is sampled for species presence and absence in N plots, sample-based rarefaction generates the expected number of accumulated species as the number of sampled plots increases from 1 to N. Accordingly, the question for this study is: can we construct a 'functional rarefaction curve' that summarizes the expected functional dissimilarity between species when n plots are drawn at random from a larger pool of N plots? Methods: In this paper, we propose a parametric measure of functional diversity that is obtained by combining sample-based rarefaction techniques that are usually applied to species richness with Rao's quadratic diversity. For a given set of N presence/absence plots, the resulting measure summarizes the expected functional dissimilarity at an increasingly larger cumulative number of plots n ( n≤ N). Results and Conclusions: Due to its parametric nature, the proposed measure is progressively more sensitive to rare species with increasing plot number, thus rendering this measure adequate for comparing the functional diversity of species assemblages that have been sampled with variable effort. [ABSTRACT FROM AUTHOR]

Abstract: In this paper, expanding on a well-known connection between Whittaker''s beta diversity and multivariate dissimilarity coefficients, a simple test for differences in beta diversity among different sets of plots is proposed. Being based on any suitable (not necessarily symmetric) plot-to-plot dissimilarity coefficient the test is appropriate for analyzing multivariate data that are usually zero-inflated due to the large number of rare species and for which the number of variables (species) is typically larger than the number of observations (plots). To illustrate the proposed test in practice, we used data from two beech forests with different management history in Abruzzo (central Italy). [Copyright &y& Elsevier]

The aim of this paper is to investigate differences in plant species composition between managed and unmanaged forests, and to assess if these difference give rise to a higher plant diversity in the unmanaged forest. Furthermore our aim is to relate forest structure to differences in plant species composition, identifying the structural attributes more strongly related to the unmanaged forest vegetation. We compared an old-growth forest and a managed highforest in the Abruzzo Lazio and Molise National Park (Central Italy). Plant species composition and diversity, deadwood components and live structure have been analyzed. We used permutational multivariate analysis of variance to test the response of species composition to management factor; furthermore, we compared species richness and beta diversity. Redundancy analysis has been used to relate plant species abundances to structural variables; the importance of dead and living wood components has been compared through variation partitioning. Plant species composition proved to be significantly different in the two sites, and the old-growth stand showed a higher plant diversity. From a structural point of view, we found differences especially in the amount and quality of deadwood, and in the diameter class distribution. These variables are also the most important in determining the old-growth stand plant species composition according to redundancy analysis. Variation partitioning confirmed the greater importance of the deadwood variables. Our results suggest that including deadwood surveys in traditional forest inventories could help in finding forests with both structural and floristic old-growth properties to be considered in conservation programmes. The imitation of natural dynamics, through the creation of gaps avoiding deadwood removal, could be an effective strategy for restoring old-growth conditions, also in terms of plant diversity. [ABSTRACT FROM AUTHOR]

Aims: Flower ultraviolet (UV) reflectance strongly influences insects' ability to detect flowers and locate pollen/nectar sources. Although included in the TRY database, the lack of a cost‐effective standardised method of measurement hampers the availability of information on this key floral trait. Digital photography and image processing allow for a novel approach to flower UV reflectance measurement that is both accessible and accurate. Location: European semi‐natural grasslands. Methods: We used a UV‐LED light and a mirrorless camera with a filter, which represents a low‐cost equipment for narrow emission/reflection photography (350–380 nm). Flowers were photographed with two standards of known reflection, and UV reflectance values were obtained using open‐source image processing software. We measured UV reflectance for 57 plant species typical of European semi‐natural grasslands. Results: Our values substantially matched the categorical classes obtained by analogue photography available in TRY and showed a highly significant relationship with spectrophotometric measures. Conclusions: The method proposed here overcomes the one based on analogue photography and subjective visual estimates, and represents an easy and low‐cost alternative to spectrophotometry. It may promote the standardised measurement of flower UV reflectance and broaden the information of this trait globally, meeting the needs of functional ecology and trait‐based community assembly studies. [ABSTRACT FROM AUTHOR]

Aim: Exploring the biotic homogenization and diversity resistance hypotheses by assessing the effect of non‐native black locust canopy on understorey species turnover. Location: Berlin, the Venetian metropolitan area, and Rome. Methods: We modelled the zeta (ζ) expression of diversity to compare the understorey species turnover between the non‐native black locust and native woodland canopies across multiple sites and through predictors of anthropogenic pressure (road and built‐up density) and interior conditions (tree basal area and mean height). Results: In Rome, black locust showed the lowest survivability and cover and did not produce any homogenization of the understorey. In Berlin and in the Venetian metropolitan area, black locust caused understorey homogenization, although with a lower intensity in Berlin. Under black locust canopies, distance between sites and road density was more consistently associated with species turnover, across urban areas and multiple sites. Under native canopies in Berlin, factors prominently associated with species turnover were road and built‐up density and mean tree height, while in the Venetian metropolitan area it was road density. Main conclusions: Evidence in support of the biotic homogenization in contrast to the diversity resistance hypothesis varied across urban areas. Species turnover was influenced by land use patterns more evidently under native tree canopies and where the non‐native tree had higher survivability. Similar analyses in other urban areas may confirm these relationships with other types of landscapes. [ABSTRACT FROM AUTHOR]

Questions: Semi-natural habitats are threatened by shifts in management with worrying effects on multiple facets of biodiversity. We revisited sites once representing a reference for a calcareous semi-natural grassland habitat aiming to: (a) identify the drivers of taxonomic, functional and phylogenetic diversity, and habitat conservation state; (b) assess the role of characteristic and derived diversity in determining these patterns; and (c) discuss the possibility of reconciling the goals of habitat conservation and enhancement of different facets of plant diversity. Location: Seven sites along the Apennines (Italy), from Mt. Catria (43.46206° N, 12.70397° E) to Mt. Alpi (40.11768° N, 15.98341° E). Methods: For 132 revisited plots, we calculated vascular plant taxonomic, functional and phylogenetic diversity using Hill numbers, and used boosted regression trees to investigate their response to the compositional dissimilarity from historical plots, to grazing intensity and to environmental variables. We identified characteristic and derived diversity and summarized them in an index of habitat conservation state whose drivers were investigated using the same approach. Results: Plant diversity was influenced by the site, whereas the habitat conservation state responded more markedly to vegetation type. Grazing intensity, slope and soil variables drove taxonomic and functional diversity, and the habitat conservation state, with some differences in their relative importance. Phylogenetic diversity responded only partly to grazing intensity, while it showed a major response to increasing temperatures. Conclusions: Patterns and drivers of different facets of plant diversity partially differ from those of the habitat conservation state, suggesting that the management of semi-natural habitats should be carefully tailored on specific conservation objectives. Generalized actions on grazing regimes and litter removal can promote habitat conservation, whereas the outcomes of these actions for plant diversity may differ across sites. Identifying areas particularly subjected to land-use changes and/or climate warming may drive conservation actions. [ABSTRACT FROM AUTHOR]

Beta‐diversity has been repeatedly shown to decline with increasing elevation, but the causes of this pattern remain unclear, partly because they are confounded by coincident variation in alpha‐ and gamma‐diversity. We used 8795 forest vegetation‐plot records from the Czech National Phytosociological Database to compare the observed patterns of beta diversity to null‐model expectations (beta‐deviation) controlling for the effects of alpha‐ and gamma‐diversity. We tested whether β‐diversity patterns along a 1200 m elevation gradient exclusively depend on the effect of varying species pool size, or also on the variation of the magnitude of community assembly mechanisms determining the distribution of species across communities (e.g. environmental filtering, dispersal limitation). The null model we used is a novel extension of an existing null‐model designed for presence/absence data and was specifically designed to disrupt the effect of community assembly mechanisms, while retaining some key features of observed communities such as average species richness and species abundance distribution. Analyses were replicated in ten subregions with comparable elevation ranges. Beta‐diversity declined along the elevation gradient due to a decrease in gamma‐diversity, which was steeper than the decrease in alpha‐diversity. This pattern persisted after controlling for alpha‐ and gamma‐diversity variation, and the results were robust when different resampling schemes and diversity metrics were used. We conclude that in temperate forests the pattern of decreasing beta‐diversity with elevation does not exclusively depend on variation in species pool size, as has been hypothesized, but also on variation in community assembly mechanisms. The results were consistent across resampling schemes and diversity measures, thus supporting the use of vegetation‐plot databases for understanding patterns of beta‐diversity at the regional scale. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

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 m2 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/datab ases/Grass landD ivers ityEx plorer) 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. [ABSTRACT FROM AUTHOR]

Questions: Which environmental factors influence fine‐grain beta diversity of vegetation and do they vary among taxonomic groups? Location: Palaearctic biogeographic realm. Methods: We extracted 4,654 nested‐plot series with at least four different grain sizes between 0.0001 m² and 1,024 m² from the GrassPlot database, covering a wide range of different grassland and other open habitat types. We derived extensive environmental and structural information for these series. For each series and four taxonomic groups (vascular plants, bryophytes, lichens, all), we calculated the slope parameter (z‐value) of the power law species–area relationship (SAR), as a beta diversity measure. We tested whether z‐values differed among taxonomic groups and with respect to biogeographic gradients (latitude, elevation, macroclimate), ecological (site) characteristics (several stress–productivity, disturbance and heterogeneity measures, including land use) and alpha diversity (c‐value of the power law SAR). Results: Mean z‐values were highest for lichens, intermediate for vascular plants and lowest for bryophytes. Bivariate regressions of z‐values against environmental variables had rather low predictive power (mean R² = 0.07 for vascular plants, less for other taxa). For vascular plants, the strongest predictors of z‐values were herb layer cover (negative), elevation (positive), rock and stone cover (positive) and the c‐value (U‐shaped). All tested metrics related to land use (fertilization, livestock grazing, mowing, burning, decrease in naturalness) led to a decrease in z‐values. Other predictors had little or no impact on z‐values. The patterns for bryophytes, lichens and all taxa combined were similar but weaker than those for vascular plants. Conclusions: We conclude that productivity has negative and heterogeneity positive effects on z‐values, while the effect of disturbance varies depending on type and intensity. These patterns and the differences among taxonomic groups can be explained via the effects of these drivers on the mean occupancy of species, which is mathematically linked to beta diversity.We analysed fine‐grain beta diversity of grasslands and other open habitats in the Palaearctic using 4,654 nested‐plot series and the z‐values of power‐law species–area relationships. Mean z‐values were highest for lichens, intermediate for vascular plants and lowest for bryophytes. The strongest environmental predictors of z‐values were herb layer cover (negative), elevation (positive), rock and stone cover (positive) and the c‐value (U‐shaped). In a conceptual figure, we summarize the resulting hypotheses how different predictors could influence z‐values via mean occupancy. [ABSTRACT FROM AUTHOR]

Aims: Primary forests are critical for forest biodiversity and provide key ecosystem services. In Europe, these forests are particularly scarce and it is unclear whether they are sufficiently protected. Here we aim to: (a) understand whether extant primary forests are representative of the range of naturally occurring forest types, (b) identify forest types which host enough primary forest under strict protection to meet conservation targets and (c) highlight areas where restoration is needed and feasible. Location: Europe. Methods: We combined a unique geodatabase of primary forests with maps of forest cover, potential natural vegetation, biogeographic regions and protected areas to quantify the proportion of extant primary forest across Europe's forest types and to identify gaps in protection. Using spatial predictions of primary forest locations to account for underreporting of primary forests, we then highlighted areas where restoration could complement protection. Results: We found a substantial bias in primary forest distribution across forest types. Of the 54 forest types we assessed, six had no primary forest at all, and in two‐thirds of forest types, less than 1% of forest was primary. Even if generally protected, only ten forest types had more than half of their primary forests strictly protected. Protecting all documented primary forests requires expanding the protected area networks by 1,132 km2 (19,194 km2 when including also predicted primary forests). Encouragingly, large areas of non‐primary forest existed inside protected areas for most types, thus presenting restoration opportunities. Main conclusion: Europe's primary forests are in a perilous state, as also acknowledged by EU's "Biodiversity Strategy for 2030." Yet, there are considerable opportunities for ensuring better protection and restoring primary forest structure, composition and functioning, at least partially. We advocate integrated policy reforms that explicitly account for the irreplaceable nature of primary forests and ramp up protection and restoration efforts alike. [ABSTRACT FROM AUTHOR]

Palaearctic grasslands are diverse and dynamic ecosystems that are in the focus of ecology, conservation biology and agronomy. This special issue is dedicated to the biodiversity and conservation issues of Palaearctic grasslands and was initiated by the Eurasian Dry Grassland Group members attending the 14th Eurasian Dry Grassland Conference (EDGC) at Sulmona, Italy in 2018. The papers in this special issue cover a wide range of grassland ecosystems from mountain dry grasslands to lowland loess grasslands, feathergrass steppes and wet grasslands, and focus on the biodiversity values and conservation issues of Palaearctic grasslands. We believe that this compilation will contribute to a better understanding of the ecology of grasslands and support their more effective conservation. [ABSTRACT FROM AUTHOR]

Questions: What is the functional trait variation of European temperate grasslands and how does this reflect global patterns of plant form and function? Do habitat specialists show trait differentiation across habitat types? Location: Europe. Methods: We compiled 18 regeneration and non‐regeneration traits for a continental species pool consisting of 645 species frequent in five grassland types. These grassland types are widely distributed in Europe but differentiated by altitude, soil bedrock and traditional long‐term management and disturbance regimes. We evaluated the multivariate trait space of this entire species pool and compared multi‐trait variation and mean trait values of habitat specialists grouped by grassland type. Results: The first dimension of the trait space accounted for 23% of variation and reflected a gradient between fast‐growing and slow‐growing plants. Plant height and SLA contributed to both the first and second ordination axes. Regeneration traits mainly contributed to the second and following dimensions to explain 56% of variation across the first five axes. Habitat specialists showed functional differences between grassland types mainly through non‐regeneration traits. Conclusions: The trait spectrum of plants dominating European temperate grasslands is primarily explained by growth strategies which are analogous to the trait variation observed at the global scale, and secondly by regeneration strategies. Functional differentiation of habitat specialists across grassland types is mainly related to environmental filtering linked with altitude and disturbance. This filtering pattern is mainly observed in non‐regeneration traits, while most regeneration traits demonstrate multiple strategies within the same habitat type. [ABSTRACT FROM AUTHOR]

Policies to mitigate climate change and biodiversity loss often assume that protecting carbon‐rich forests provides co‐benefits in terms of biodiversity, due to the spatial congruence of carbon stocks and biodiversity at biogeographic scales. However, it remains unclear whether this holds at the scales relevant for management, and particularly large knowledge gaps exist for temperate forests and for taxa other than trees. We built a comprehensive dataset of Central European temperate forest structure and multi‐taxonomic diversity (beetles, birds, bryophytes, fungi, lichens, and plants) across 352 plots. We used Boosted Regression Trees (BRTs) to assess the relationship between above‐ground live carbon stocks and (a) taxon‐specific richness, (b) a unified multidiversity index. We used Threshold Indicator Taxa ANalysis to explore individual species' responses to changing above‐ground carbon stocks and to detect change‐points in species composition along the carbon‐stock gradient. Our results reveal an overall weak and highly variable relationship between richness and carbon stock at the stand scale, both for individual taxonomic groups and for multidiversity. Similarly, the proportion of win‐win and trade‐off species (i.e., species favored or disadvantaged by increasing carbon stock, respectively) varied substantially across taxa. Win‐win species gradually replaced trade‐off species with increasing carbon, without clear thresholds along the above‐ground carbon gradient, suggesting that community‐level surrogates (e.g., richness) might fail to detect critical changes in biodiversity. Collectively, our analyses highlight that leveraging co‐benefits between carbon and biodiversity in temperate forest may require stand‐scale management that prioritizes either biodiversity or carbon in order to maximize co‐benefits at broader scales. Importantly, this contrasts with tropical forests, where climate and biodiversity objectives can be integrated at the stand scale, thus highlighting the need for context‐specificity when managing for multiple objectives. Accounting for critical change‐points of target taxa can help to deal with this specificity, by defining a safe operating space to manipulate carbon while avoiding biodiversity losses. Can managing forests for storing carbon jointly achieve biodiversity conservation and climate change mitigation goals? Global and regional environmental policies often assume co‐benefits between carbon and biodiversity, but it is unclear whether this assumption holds at fine scales relevant for management. By modelling the carbon–biodiversity relationship for a broad range of taxa in temperate forests, we found that biodiversity and carbon stocks are not well‐aligned spatially at stand‐scale, and their relationship varied substantially across taxa. This suggests that forest management aimed at storing carbon may benefit some taxa but harm others, highlighting that broad‐scale co‐benefits may break down at fine scales. [ABSTRACT FROM AUTHOR]

Monitoring semi-natural grasslands is difficult and costly because they occur in highly dynamic and extremely complex habitat mosaics. We combined the use of a low-cost unmanned aerial vehicle (UAV) and Structure from Motion (SfM) photogrammetry to acquire high spatial resolution (∼1.5 cm pixel) RGB imagery. After image classification through Object-Based Image Analysis (OBIA), we accurately were able to distinguish three semi-natural grassland types, one of which is a habitat of conservation concern. The use of orthomosaics, digital elevation models (DEMs), and canopy height models (CHMs) yielded excellent overall classification accuracies (>89%) assessed through both remotely validated and ground-truthed points. We identified two layers of woody vegetation with a user's (UA) and producer's (PA) accuracies >73% and three grassland types: closed grassland (UA = 94%; PA = 97%), open grassland habitat (UA = 97%; PA = 93%) and open grasslands with soil erosion (UA = 96%; PA = 98%). The grassland types differed substantially in the cover of vegetation, rocks, stones, and bare soil measured in the field, as well as in the number and relative cover of the habitat diagnostic species. The proposed methodology is highly promising for mapping and monitoring semi-natural grassland of conservation concern in support of tailored management actions. [Display omitted] • Fine-scale mapping identifies three different semi-natural grassland classes. • High-resolution imagery and dense point clouds improve vegetation classification. • A useful tool for monitoring habitats of conservation concern. • Cost-effective and time-saving tool for frequent monitoring programs. [ABSTRACT FROM AUTHOR]

Assessing the contribution of forest soils to carbon (C) and nutrient dynamics is extremely complex due to the high spatial variability of soil properties, at fine to very broad scales. Improving our understanding of soil variability is necessary to scale up sample-based inventory data for the estimation of regional contribution of forest soils to C dynamics. We analyzed the topsoil spatial variability of organic carbon (OC) and matter (OM), nitrogen (N), C/N ratio, texture, and pH in 11 southern European old-growth beech stands. Our aim was to assess the within- and among-stand topsoil variability, and to determine the drivers underlying this variation. For each stand, we sampled the topsoil in 25 quadrats in a 1-ha regular grid where we recorded overstorey structural and compositional attributes, deadwood quantity and quality, microtopography and site conditions. Soil parameters varied highly at both scales: When considering all the topsoil properties together through multivariate analysis, most of the variability occurred at the within-stand scale (mostly due to shifts in organic matter content, C/N, and litter depth); the opposite was true when considering soil properties singularly, with pH, soil texture, and N concentration varying greatly among stands. Most of the among-stand variability depended on climate and on the direct and indirect effects of parent material. Fine-scale variation, instead, depended mainly on overstorey composition and microtopographical variation. Surprisingly, we found no direct influence of overstorey structure on topsoil parameters, likely because soil parameters respond to changes in stand structural features only after a substantial time lag. [ABSTRACT FROM AUTHOR]

In Europe, the conservation of extensively grazed semi-natural grasslands is addressed by agricultural policies whose effectiveness is questioned. We studied sub-xerophilous Bromus erectus semi-natural grasslands to analyse the interactions among: i) agri-environmental payments, ii) grazing regimes, iii) environmental conditions, iv) habitat conservation state, and v) forage yield and quality. We sampled 98 plots across 19 farms and unmanaged control areas in five regions encompassing Italy and Switzerland. We fitted two piecewise structural equation models (SEM) to infer direct and indirect effects of agri-environmental payments, grazing regimes and environmental conditions on proxies of habitat conservation state, (i.e., the number and cover of diagnostic species), and forage yield and quality (i.e., specific leaf area - SLA, leaf dry-matter content - LDMC, sward height and pastoral value). Agri-environmental payments contributed to maintain grazing management and in turn to preserve the habitat biodiversity and functions. Payments did not affect stocking rates, but determined a more even distribution of grazing intensity, with positive effects on habitat conservation state and negative outcomes for LDMC. Conversely, LDMC increased with stocking rates. Among environmental condition, elevation and soil carbonates content had a positive effect on the habitat conservation state, while slope exerted only indirect effects on forage quality and diagnostic species by reducing fine-scale grazing intensity. Overall, the effectiveness of payments largely depended on the scale of measures' implementation. Farm-level grazing contracts and periodic field monitoring would allow to influence the fine-scale grazing intensity and to implement a result-oriented approach towards the objectives of the post-2020 CAP. • Payments do not influence stocking rates, but fine-scale grazing intensity. • Fine-scale grazing intensity affects habitat conservation and forage provision. • Stocking rates affect forage yield and quality, but not habitat conservation state. • Elevation, soil carbonates and slope influence habitat conservation state. • Payments should include farm specific grazing contracts and habitat monitoring. [ABSTRACT FROM AUTHOR]

We performed our study in the Dry Puna of the southern Peruvian Andes. Through a comparative approach we aimed to assess the effects of the two management systems, low grazing pressure by wild camelids vs. high grazing pressure by domestic livestock and periodic burning. Our general hypothesis was that the traditional high disturbance regime affects the dry Puna species diversity and composition through modifications of the magnitude of plant–plant-interactions and changes of the community structure due to shifts in species dominance. In 40 plots of 10 × 10 m, the cover value of each species was recorded and the species richness, floristic diversity, and community similarity of each treatment were compared. For each disturbance regime, differences of soil features (organic matter, carbon/nitrogen ratio, and potassium content) were tested. To evaluate plant–plant interactions, 4 linear transect divided into 500 plots of 10 × 10 cm were laid out and co-occurrence analysis was performed. We found that different disturbance regimes were associated with differences in the floristic composition, and that the high disturbance condition had lower species diversity and evenness. A decrease of tall species such as Festuca orthophylla and increase of dwarf and spiny Tetraglochin cristatum shrubs was observed as well. In addition, different disturbance intensities caused differences in the functional composition of the plant communities, since species with avoidance strategies are selected by high grazing pressure. High disturbance intensity was also associated to differences of soil features and to different clumped spatial structure of the dry Puna. Our results indicate also that: positive interactions are often species-specific mainly depending on the features of nurse and beneficiary species; the importance of positive interaction is higher at low grazing pressure than at high disturbance intensity; the magnitude and direction of the herbivory-mediated facilitation processes may be traced back to the grazing pressure of wild camelids. [ABSTRACT FROM AUTHOR]