Summary Species distribution models (SDMs) have become a standard tool in ecology and applied conservation biology. Modelling rare and threatened species is particularly important for conservation purposes. However, modelling rare species is difficult because the combination of few occurrences and many predictor variables easily leads to model overfitting. A new strategy using ensembles of small models was recently developed in an attempt to overcome this limitation of rare species modelling and has been tested successfully for only a single species so far. Here, we aim to test the approach more comprehensively on a large number of species including a transferability assessment. For each species, numerous small (here bivariate) models were calibrated, evaluated and averaged to an ensemble weighted by AUC scores. These ‘ensembles of small models’ (ESMs) were compared to standard SDMs using three commonly used modelling techniques (GLM, GBM and Maxent) and their ensemble prediction. We tested 107 rare and under‐sampled plant species of conservation concern in Switzerland. We show that ESMs performed significantly better than standard SDMs. The rarer the species, the more pronounced the effects were. ESMs were also superior to standard SDMs and their ensemble when they were evaluated using a transferability assessment. By averaging simple small models to an ensemble, ESMs avoid overfitting without losing explanatory power through reducing the number of predictor variables. They further improve the reliability of species distribution models, especially for rare species, and thus help to overcome limitations of modelling rare species.

Ensembles of Small Models (ESM) represent a novel strategy for species distribution modelling with few observations. ESMs are built by calibrating many small models and then averaging them into an ensemble model where the small models are weighted by their cross‐validated scores of predictive performance. In a previous paper (Breiner, Guisan, Bergamini, & Nobis, Methods in Ecology and Evolution, 6, 1210–1218, 2015), we reported two major findings. First, ESMs proved largely superior to standard models in terms of model performance and transferability. Second, ESMs including different modelling techniques did not clearly improve model performance compared to single‐technique ESMs. However, ESMs often require a large computation effort, which can become problematic when modelling large numbers of species. Given the appealing new perspectives offered by ESMs, it is especially important to investigate if some techniques yield increased performance while saving computation time and thus could be predominantly used for building ESMs. Here, we present results from a reanalysis of a subset of the data used in Breiner et al. (2015). More specifically, we ran ESMs: (1) fitted with 10 modelling techniques separately (in Breiner et al., 2015 we used only three techniques); and (2) using various parameter options for each modelling technique (i.e., model tuning). We show that ESMs vary in model performance and computation time across techniques, and some techniques are advantageous in terms of optimizing model performance and computation time (i.e., GLM, CTA and ANN). Including one of these modelling techniques could thus optimize computation time compared to using more computing‐intensive techniques like GBM. Next, we show that parameter tuning can improve performance and transferability of ESMs, but often at the cost of computation time. Parameter tuning could therefore be used when computing resources are not a limiting factor. These findings help improve the applicability and performance of ESMs when applied to large numbers of species.

Aims Phytosociological classification of fen vegetation (Scheuchzerio palustris‐Caricetea fuscae class) differs among European countries. Here we propose a unified vegetation classification of European fens at the alliance level, provide unequivocal assignment rules for individual vegetation plots, identify diagnostic species of fen alliances, and map their distribution. Location Europe, western Siberia and SE Greenland. Methods 29 049 vegetation‐plot records of fens were selected from databases using a list of specialist fen species. Formal definitions of alliances were created using the presence, absence and abundance of Cocktail‐based species groups and indicator species. DCA visualized the similarities among the alliances in an ordination space. The ISOPAM classification algorithm was applied to regional subsets with homogeneous plot size to check whether the classification based on formal definitions matches the results of unsupervised classifications. Results The following alliances were defined: Caricion viridulo‐trinervis (sub‐halophytic Atlantic dune‐slack fens), Caricion davallianae (temperate calcareous fens), Caricion atrofusco‐saxatilis (arcto‐alpine calcareous fens), Stygio‐Caricion limosae (boreal topogenic brown‐moss fens), Sphagno warnstorfii‐Tomentypnion nitentis (Sphagnum‐brown‐moss rich fens), Saxifrago‐Tomentypnion (continental to boreo‐continental nitrogen‐limited brown‐moss rich fens), Narthecion scardici (alpine fens with Balkan endemics), Caricion stantis (arctic brown‐moss rich fens), Anagallido tenellae‐Juncion bulbosi (Ibero‐Atlantic moderately rich fens), Drepanocladion exannulati (arcto‐boreal‐alpine non‐calcareous fens), Caricion fuscae (temperate moderately rich fens), Sphagno‐Caricion canescentis (poor fens) and Scheuchzerion palustris (dystrophic hollows). The main variation in the species composition of European fens reflected site chemistry (pH, mineral richness) and sorted the plots from calcareous and extremely rich fens, through rich and moderately rich fens, to poor fens and dystrophic hollows. ISOPAM classified regional subsets according to this gradient, supporting the ecological meaningfulness of this classification concept on both the regional and continental scale. Geographic/macroclimatic variation was reflected in the second most important gradient. Conclusions The pan‐European classification of fen vegetation was proposed and supported by the data for the first time. Formal definitions developed here allow consistent and unequivocal assignment of individual vegetation plots to fen alliances at the continental scale.

Changing land use has a major impact on lichen diversity. This study attempts to identify patterns or trends of lichen functional groups along a land use gradient, ranging from natural forests to open agricultural landscape. In eight countries, covering six main European biogeographic regions, lichen vegetation was assessed according to a standardized scheme. Data on reproductive, vegetative and ecological traits was compiled and relative species richness for all classes of all traits calculated. Relationships between the land use gradient and relative species richness of trait classes were analysed. Open and intensively managed landscapes harbour more fertile species while sterile species are relatively more important in forests. This finding is also supported by analyses of different classes of dispersal propagules. The importance of species with the principal photobiont Trebouxia s.l. increases linearly with intensification of land use. A converse pattern is revealed by species with Trentepohlia. Concerning substratum specialization only generalists show an effect along the land use intensity gradient. Their relative species richness decreases from landscapes dominated by forests to open agricultural landscape. A considerable decline in the rare lichen species richness as a result of land intensification is predicted.

Fig. 2. Spatial coverage of GrassPlot data from Morocco to Japan. Currently, the majority comes from sub-Mediterranean to hemiboreal Europe (black = multi-scale plots, grey = other plots). Current content v. 1.00 (January 2018) • 126 datasets • 198 data owners • 36 countries • 168,997 plots, among them 14,064 with data also for non-vascular plants • 66,000 0.01-m² plots, 17,206 1-m² plots, 5,520 10-(or 9-) m² plots, 2,545 100-m² plots • 2,797 nested-plot series (with at least 4 grain sizes)

Aim Information on change in species’ environmental preferences (i.e., niche) is currently not included in IUCN Red List criteria, although such information is key for assessing whether species not only lose geographic range but also lose part of their realized niche. Here, using niche size quantification and niche‐based species distribution models (SDMs), we test whether realized niche size and predicted potential range size provide additional information compared with the standard IUCN scores. Location Switzerland, national scale. Methods We simulated randomly, spatially directed, and ecologically directed local extinction events of varying magnitudes (10%, 30%, and 50% of occurrences). For a set of 148 representative vascular plant species, we tested how accurately the geographic versus niche measures pictured these extinction scenarios respectively. Results We found that changes in niche size often corresponded to changes in geographic space. However, there was considerable variation and, for many species, changes in geographic and in niche space delivered complementary information. IUCN criteria based on spatial projections of SDMs did not capture extinction events in most cases and often increased the modelled range size, even when up to 50% of the occurrences were removed by simulated extinction events. Main conclusion Our findings demonstrate that changes in niche size can provide valuable additional information and could be used more systematically to complement changes in range size for Red List assessments. In turn, change in SDM‐predicted range size was not a good surrogate for classical extent of occurrence and area of occupancy criteria and should be used with caution. Further research is needed to assess whether and how spatial predictions of SDMs may be used to appropriately complement current IUCN criteria and to test whether our findings apply to other organisms and other spatial extents.

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Many ecosystems of high conservation value have been shaped by human impacts over centuries. Today,traditional management of semi-natural habitats is a common conservation measure in Europe. However, despite traditional management, habitat remnants may still loose specialist species due to surrounding land-use change or atmospheric nitrogen deposition. To detect trends in species density (2-m2 plotscale) and habitat quality in calcareous fens in the pre-Alps of Switzerland, we surveyed 36 traditionally managed fens in 1995/97 and again in 2005/06 (five plots per fen). The fens occurred at three altitudinal levels (800-1000, 1000-1200, 1200-1400 m asl) and were either extensively grazed or mown once a year. Despite these traditional management regimes, species density of fen specialists and of all bryophytes decreased during this decade (vascular plant specialists: -9.4%, bryophytespecialists: -14.9%, all bryophytes: -5.7%). Management had no effect on the number of Red-List species and habitat specialists of vascular plants per plot. However, bryophyte species density was more strongly reduced in grazed fens. Species density of vascular plant generalists increased between the two surveys(+8.2%) but not of bryophytes. Among vascular plants, Red-List species decreased from 1.01 to 0.78 species per plot. Furthermore, between the two surveys aboveground plant biomass, mean plant-community indicator values for nutrients and species density of nutrient indicators increased, whereas mean plant indicator values for soil moisture, light and peat, and species density for peat indicators, decreased. We attribute these changes and the loss of specialist species over the past decade mainly to land-use change in the surrounding area and to nutrient inputs. Thus, despite traditional management, calcareous fens in the pre-Alps suffer from ongoing habitat deterioration and endangered plant species remain threatened. For their long-term protection, we suggest to reduce nutrient inputs and, where necessary to restore hydrology and adjust grazing management. plots per fen). The fens occurred at three altitudinal levels (800-1000, 1000-1200, 1200-1400 m 28 a.s.l.) and were either extensively grazed or mown once a year. Despite these traditional 29 management regimes, species density of fen specialists and of all bryophytes decreased during 30 this decade (vascular plant specialists: -9.4%, bryophyte specialists: -14.9%, all bryophytes: -31 5.7%). Management had no effect on the number of Red-List species and habitat specialists of 32 vascular plants per plot. However, bryophyte species density was more strongly reduced in 33 grazed fens. Species density of vascular plant generalists increased between the two surveys 34 (+8.2%) but not of bryophytes. Among vascular plants, Red-List species decreased from 1.01 to 35 0.78 species per plot. Furthermore, between the two surveys aboveground plant biomass, mean 36 plant-community indicator values for nutrients and species density of nutrient indicators 37 increased, whereas mean pl...