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Secondary dry grasslands in Europe can host high levels of vascular plant richness at small spatial scales. However, in Southern Europe their biodiversity patterns are largely unexplored. In this work, we aim at: (i) estimating plant species richness patterns at very fine scales in montane dry grasslands, on limestone bedrock, in Abruzzo Lazio and Molise National Park (Central Apennines, Italy); (ii) assessing the most important physical and edaphic drivers of biodiversity patterns at multiple plot sizes. We used randomly placed nested-plot series where we measured alpha-diversity at three different plot sizes (1 m2, 0.1 m2 and 0.01 m2) and within-plot beta-diversity (as expressed by the slope of the species-area curve across plot sizes). Variable selection was performed by means of Random Forests. Relationships between selected variables and diversity measures were then assessed using Regression Trees, Linear and Generalized Linear Models. Overall, results pointed to topographically-controlled edaphic factors (soil pH and silt fraction) as the main drivers positively influencing alpha-diversity at all spatial scales, with a positive effect of rock cover and slope inclination at smaller spatial grains. Beta-diversity was positively influenced by rock cover. We suggest that high-pH, steep and/or rocky sites feature higher species richness because they lack competitive grass species. Our results are in agreement with previous works underlining the importance of less productive habitats for the conservation of secondary grassland biodiversity.

Intraspecific trait variation (ITV), based on available genetic diversity, is one of the major means plant populations can respond to environmental variability. The study of functional trait variation and diversity has become popular in ecological research, for example, as a proxy for plant performance influencing fitness. Up to now, it is unclear which aspects of intraspecific functional trait variation (iFDCV) can be attributed to the environment or genetics under natural conditions. Here, we examined 260 individuals from 13 locations of the rare (semi‐)dry calcareous grassland species Trifolium montanum L. in terms of iFDCV, within‐habitat heterogeneity, and genetic diversity. The iFDCV was assessed by measuring functional traits (releasing height, biomass, leaf area, specific leaf area, leaf dry matter content, Fv/Fm, performance index, stomatal pore surface, and stomatal pore area index). Abiotic within‐habitat heterogeneity was derived from altitude, slope exposure, slope, leaf area index, soil depth, and further soil factors. Based on microsatellites, we calculated expected heterozygosity (He) because it best‐explained, among other indices, iFDCV. We performed multiple linear regression models quantifying relationships among iFDCV, abiotic within‐habitat heterogeneity and genetic diversity, and also between separate functional traits and abiotic within‐habitat heterogeneity or genetic diversity. We found that abiotic within‐habitat heterogeneity influenced iFDCV twice as strong compared to genetic diversity. Both aspects together explained 77% of variation in iFDCV (Radj2 = .77, F2, 10 = 21.66, p < .001). The majority of functional traits (releasing height, biomass, specific leaf area, leaf dry matter content, Fv/Fm, and performance index) were related to abiotic habitat conditions indicating responses to environmental heterogeneity. In contrast, only morphology‐related functional traits (releasing height, biomass, and leaf area) were related to genetics. Our results suggest that both within‐habitat heterogeneity and genetic diversity affect iFDCV and are thus crucial to consider when aiming to understand or predict changes of plant species performance under changing environmental conditions. Up to now, it is unclear which aspect of intraspecific functional trait variation (ITV) can be attributed to the environment or to genetics under natural and present environmental conditions. Here, we used data from an extensive field study (260 individuals from 13 locations) on the (semi-)dry calcareous grassland species Trifolium montanum (mountain clover). We demonstrated that abiotic within-habitat heterogeneity had twice as much impact as genetic diversity on ITV explaining together 77% of variation.

Question: Can vegetation relevé databases be used to analyse species losses and gains in specific vegetation types in Germany over time? Does the type of response (increase or decline in relative frequency) conform to observed large-scale environmental trends in the last decades? Location: Germany. Exploring the German Vegetation Reference Database Halle (GVRD) that was established for forest and grassland vegetation within the framework of German Biodiversity Exploratories. Methods: Use of generalized linear models (GLMs) for testing changes in temporal frequency of plant taxa in a semi-dry grassland data set (Mesobromion) and a beech forest data set (Fagion). Data were either aggregated by year, decade or by a balanced re-sampling approach. Interpretation of the observed changes was based on species traits. Results: In both data sets significant temporal changes were observed, although the frequency of the majority of species remained unchanged. In both data sets, species with a temporal increase in frequency had higher Ellenberg N and F indicator values, compared to species that decreased, thus indicating effects of widespread atmospheric nitrogen deposition. In the forest data set, the observed increase in recruitment of deciduous trees pointed to a change in management, while trends in the grassland data set suggested use abandonment, as seen in an increased frequency of woody species. Conclusion: We demonstrate that vegetation databases represent very valuable resources for analysis of temporal changes in species frequencies. GLMs proved their value in detecting these trends, as also shown by the interpretability of model results with species traits. In contrast, the method of aggregation or re-sampling had little influence on the general outcome of analyses.

Questions Has plant species richness in semi‐natural grasslands changed over recent decades? Do the temporal trends of habitat specialists differ from those of habitat generalists? Has there been a homogenization of the grassland vegetation? Location Different regions in Germany and the UK. Methods We conducted a formal meta‐analysis of re‐survey vegetation studies of semi‐natural grasslands. In total, 23 data sets were compiled, spanning up to 75 years between the surveys, including 13 data sets from wet grasslands, six from dry grasslands and four from other grassland types. Edaphic conditions were assessed using mean Ellenberg indicator values for soil moisture, nitrogen and pH. Changes in species richness and environmental variables were evaluated using response ratios. Results In most wet grasslands, total species richness declined over time, while habitat specialists almost completely vanished. The number of species losses increased with increasing time between the surveys and were associated with a strong decrease in soil moisture and higher soil nutrient contents. Wet grasslands in nature reserves showed no such changes or even opposite trends. In dry grasslands and other grassland types, total species richness did not consistently change, but the number or proportions of habitat specialists declined. There were also considerable changes in species composition, especially in wet grasslands that often have been converted into intensively managed, highly productive meadows or pastures. We did not find a general homogenization of the vegetation in any of the grassland types. Conclusions The results document the widespread deterioration of semi‐natural grasslands, especially of those types that can easily be transformed to high production grasslands. The main causes for the loss of grassland specialists are changed management in combination with increased fertilization and nitrogen deposition. Dry grasslands are most resistant to change, but also show a long‐term trend towards an increase in more mesotrophic species. Semi‐natural grasslands are famous for their high plant species diversity. In a meta‐analysis of 23 data sets from Germany and the UK we show that the total species richness in wet grasslands has declined over time, while in dry grasslands the number of species did not consistently change. However, there was a general decrease in the proportion of habitat specialists.

Revisiting the classical Gradmann’s ‘steppe theory’ for central Europe, we examine whether the early Holocene steppe habitats survived the critical period of maximum Holocene afforestation: the mid-Holocene bottleneck. Despite the undisputable fact that afforestation was a dominant ecological factor in this period, our parallel analyses of pollen and molluscs from sedimentary sequences discovered in the dry lowland area of northern Bohemia, Czech Republic (Zahájí and Suchý potok sites, lower Ohře area) provide strong evidence for uninterrupted local occurrence of steppe grasslands throughout the Holocene. At the onset of the Neolithic agriculture, this area was covered by forest-steppe. Analogously to the present forest-steppe landscapes of eastern Europe and south-western Siberia, dry areas of northern Bohemia were dominated by open-canopy pine–birch forests that enabled continuous survival of many light-demanding plant species from the late Glacial and early Holocene to the Neolithic. Later on, anthropogenic deforestation and livestock grazing created a semi-natural steppe. Our data suggest that this secondary steppe can be viewed as a direct continuation of the late Pleistocene and early Holocene natural steppe rather than a purely cultural steppe developed only after deforestation of a continuously forested mid-Holocene landscape by humans. At the same time, we provide evidence supporting Gradmann’s ‘steppe theory’, assuming that in central Europe, Neolithic farming started in those areas that were not completely forested but contained remnants of natural steppes. This finding has important implications for the interpretation of present biodiversity patterns in central Europe.

Aim: Soil pH is considered an important driver of fine-scale plant species richness in terrestrial ecosystems. However, it is unclear to what extent this relationship is influenced by precipitation, which often directly affects both soil pH and species richness. We asked: (1) what is the relationship between fine-scale vascular plant species richness and soil pH in regions with different levels of precipitation and (2) what are the relative effects of soil pH and precipitation on species richness? Location: Dry grasslands in eight regions of northern Eurasia. Methods: Species richness and soil pH were measured in 1055 10 m × 10 m plots and precipitation values were derived from global dataseis. Relationships between variables were explored using general linear models, mixed-effect models and partial regressions. Variation partitioning was used to assess the relative effect of each predictor on species richness. Results: In wetter regions, soil pH range was broader, mean species richness was higher and the richness-pH relationship was unimodal. In drier regions, mean soil pH was higher and its range narrower, species richness was on average lower and less variable, and the richness-pH relationship was negative or absent. The richness-pH relationship persisted after controlling for the effect of precipitation, but precipitation, uniquely or together with soil pH, explained more variation in species richness in most regions than did pH alone. Main conclusions: The relationship between plant species richness and soil pH in dry grasslands changes from unimodal, through negative, to none with decreasing regional precipitation in Eurasia. However, it seems that the species richness-soil pH relationship in dry grasslands over broad areas is substantially influenced and confounded by precipitation either indirectly, by shortening and shifting the pH gradient, or directly, by decreasing the negative effects of drought stress on richness.

The abandonment of agricultural lands triggers a secondary succession of plant species which implies important changes in soil quality. Annual Mediterranean grasslands are known to be persistent on abandoned agriculture lands in the western Mediterranean. We used plant taxonomic and functional approaches to determine the role of Mediterranean grasslands as an indicator of changes in ecosystem biodiversity and functionality. We tested the hypothesis that Mediterranean grasslands are a suitable model for monitoring biodiversity and soil fertility in a secondary succession. Soil and vegetation features on 21 permanent plots were monitored in 2016 and 2020. Numerical classifications based on floristic composition showed two different plant communities independently of the sampling year: early-stage grasslands in the first post-abandonment decade and late-stage grasslands after the first post-abandonment decade. Generalized linear model and redundancy analysis also revealed differences in growth forms, functional traits and soil functionality between communities. Late-stage grasslands was characterized by enriched bryophyte coverage and an impoverishment in hemicryptophytes and plant latex segregators growing on soils with a higher hydrolase enzyme activity and TOC content compared to early-stage grassland. Our results suggest that annual Mediterranean grasslands growing on siliceous soils denoting a mature-stage succession, and floristically characterized by the symbiont plant with Ascomycota, Tuberaria guttata, and a high bryophyte cover, are worthy of recognition for conservation.

Plants inhabiting open landscapes are often dispersed by ungulates and are expected to be adapted to this type of dispersal through their seed traits. To find which traits help seeds survive the passage through digestion of wild ungulates, we conducted a comprehensive feeding experiment with almost forty species of plants and three species of ungulates. We fed specified numbers of seeds to the animals, collected the dung, and germinated the dung content. We explored whether seed morphological traits and seed nutrient contents are good predictors of seed survival after passage through the ungulate digestive system. We also tested how the seed survival differed after the passage through different ungulate species. To find answers, we used GLMM with beta-binomial distribution and animal and plant species as random factor, respectively. We found that species survival and germination success were negatively correlated to seed elongation and the thickness of the seed coat. Even though phylogenetically correct GLMM did not yield significant results, when we tested species from commonly represented families, separately (legumes and grasses compared to all other species) different traits had statistically significant effects. In the case of seed elongation, the effect changed direction from negative to positive when legumes and grasses were left out. Our results suggest that seed traits enabling species survival after passage through the digestive tract are strongly phylogenetically conserved and different groups of plants evolved different ways of adapting to grazing pressure and utilize it for dispersal.

Aims This study investigates how precipitation, temperature and seasonality (as a proxy of plant productivity) affect the temporal and spatial variability of soil CO 2 efflux in two dry semiarid grasslands with different degrees of land degradation. Methods We measured soil CO 2 efflux over four years under plant, biological soil crust and bare soil patches and estimated annual soil carbon losses in both, a natural and a degraded grassland, by means of generalised additive mixed models considering temporal autocorrelation in the data. Results Soil CO 2 efflux ranged from 0.08 to 3.70 and from 0.10 to 3.01 μmol CΟ 2 m −2  s −1 in the natural and degraded grasslands, respectively. Daily soil CO 2 efflux was mostly affected by moisture in the degraded grassland (25.4%), while in the natural grassland was affected jointly by seasonality, temperature and moisture (27.5%). Overall, the highest soil carbon fluxes were measured in soils covered by biological soil crusts (1.24 ± 0.02 and 1.10 ± 0.02) and the lowest in bare soils (1.11 ± 0.02 and 0.82 ± 0.02 μmol CΟ 2 −2  s −1 ) in the natural and degraded sites, respectively. Cumulative soil carbon fluxes were mainly driven by temperature and previous precipitation (over three months). The highest soil carbon losses were estimated in the driest year (2009) and the lowest in the wettest (2010) with almost twice the amount of rainfall. The main difference between these years was the timing of the events that mostly occurred in the moments of maximum plant activity with optimum temperatures in spring in the dry year. Conclusions Changes in precipitation patterns will affect soil carbon fluxes more than rainfall amount, particularly in degraded grasslands. Therefore, considering all climate drivers together with plant activity is essential to predict how climate change will affect soil biological processes in drylands.

Aim Species on islands are at high risk of extinction due to environmental changes, including global warming, land‐use alterations and invasions. At local scales, extinctions can be offset by strategies promoting in situ persistence. We explored how persistence‐related traits of plants—that is, linked to belowground resource conservation, growth, size and longevity—on edaphic islands respond to variation in insularity and the environment (soil and microclimate), including intraspecific variability, which is rarely considered in functional island biogeography. We hypothesised that plants facing strong insularity and harsh soil conditions are characterised by enhanced persistence abilities. Location Shallow‐soil temperate dry grasslands on granite outcrops, Central Europe. Methods We focussed on edaphic island specialist species belonging to different life histories, namely clonal and non‐clonal perennial plants. We used linear and linear mixed‐effect models to examine intra‐ and interspecific trait patterns versus variation in insularity, soil and microclimate. Results Insularity tended to promote smaller plants (non‐clonal species) and belowground resource‐conservative strategies (both clonal and non‐clonal species), increasing the likelihood of local persistence. Soil also contributed largely to explaining persistence‐related trait patterns: plants growing in harsh soil conditions tended to be resource conservative. Clonal species are distinguished by highly consistent responses to variation in insularity and soil conditions, whereas non‐clonal plants showed distinct species‐specific responses. Main conclusions Our findings have important implications for the conservation biogeography of edaphic island plant specialists. Clonal species may be susceptible to local extinction should insularity or soil conditions vary, for example, due to abrupt changes in the geographical setting (e.g. habitat loss) or local environmental factors (e.g. N‐deposition). Non‐clonal species may instead face environmental changes differently; some will go extinct, whereas others will survive, depending on the prevailing abiotic pressures. This seems to challenge previous views that predicted clonal species to be the winners and non‐clonal species the losers against local extinction.