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Human-induced nitrogen (N) and phosphorus (P) enrichment have profound effects on grassland net primary production (NPP) and species richness. However, a comprehensive understanding of the relative contribution of N vs. P addition and their interaction on grassland NPP increase and species loss remains elusive. We compiled data from 80 field manipulative studies and conducted a meta-analysis (2107 observations world-wide) to evaluate the individual and combined effects of N and P addition on grassland NPP and species richness. We found that both N addition and P addition significantly enhanced grassland above-ground NPP (ANPP; 33.2% and 14.2%, respectively), but did not affect total NPP, below-ground NPP (BNPP), and species evenness. Species richness significantly decreased with N addition (11.7%; by decreasing forbs) probably due to strong decreased soil pH, but not with P addition. The combined effects of N and P addition were generally stronger than the individual effects of N or P addition, and we found the synergistic effects on ANPP, and additive effects on total NPP, BNPP, species richness, and evenness within the combinations of N and P addition. In addition, N and P addition effects were strongly affected by moderator variables (e.g. climate and fertilization type, duration and amount of fertilizer addition). These results demonstrate a higher relative contribution of N than P addition to grassland NPP increase and species loss, although the effects varied across climate and fertilization types. The existing data also reveals that more long-term (≥5 years) experimental studies that combine N and P and test multifactor effects in different climate zones (particularly in boreal grasslands) are needed to provide a more solid basis for forecasting grassland community response and C sequestration response to nutrient enrichment at the global scale.

Despite the ecological significance of ericoid mycorrhizal fungi, little is known about the abiotic and biotic factors driving their diversity and community composition. To determine the relative importance of abiotic and biotic filtering in structuring ericoid mycorrhizal fungal communities, we established 156 sampling plots in two highly contrasting environments but dominated by the same Ericaceae plant species: waterlogged bogs and dry heathlands. Plots were located across 25 bogs and 27 dry heathlands in seven European countries covering a gradient in nitrogen deposition and phosphorus availability. Putatively ericoid mycorrhizal fungal communities in the roots of 10 different Ericaceae species were characterized using high-throughput amplicon sequencing. Variation in ericoid mycorrhizal fungal communities was attributed to both habitat and soil variables on the one hand and host plant identity on the other. Communities differed significantly between bogs and heathlands and, in a given habitat, communities differed significantly among host plant species. Fungal richness was negatively related to nitrogen deposition in bogs and phosphorus availability in bogs and heathlands. Our results demonstrate that both abiotic and biotic filtering shapes ericoid mycorrhizal fungal communities and advocate an environmental policy minimizing excess nutrient input in these nutrient-poor ecosystems to avoid loss of ericoid mycorrhizal fungal taxa.

Background Ecosystem restoration is as a critical tool to counteract the decline of biodiversity and recover vital ecosystem services. Restoration efforts, however, often fall short of meeting their goals. Although functionally important levels of biodiversity can significantly contribute to the outcome of ecosystem restoration, they are often overlooked. One such important facet of biodiversity is within-species genetic diversity, which is fundamental to population fitness and adaptation to environmental change. Also the diversity of arbuscular mycorrhizal fungi (AMF), obligate root symbionts that regulate nutrient and carbon cycles, potentially plays a vital role in mediating ecosystem restoration outcome. In this study, we investigated the relative contribution of intraspecific population genetic diversity, AMF diversity, and their interaction, to population recovery of Succisa pratensis, a key species of nutrient poor semi natural grasslands. We genotyped 180 individuals from 12 populations of S. pratensis and characterized AMF composition in their roots, using microsatellite markers and next generation amplicon sequencing, respectively. We also investigated whether the genetic makeup of the host plant species can structure the composition of root-inhabiting AMF communities. Results Our analysis revealed that population allelic richness was strongly positively correlated to relative population growth, whereas AMF richness and its interaction with population genetic diversity did not significantly contribute. The variation partitioning analysis showed that, after accounting for soil and spatial variables, the plant genetic makeup explained a small but significant part of the unique variation in AMF communities. Conclusions Our results confirm that population genetic diversity can contribute to population recovery, highlighting the importance of within-species genetic diversity for the success of restoration. We could not find evidence, however, that population recovery benefits from the presence of more diverse AMF communities. Our analysis also showed that the genetic makeup of the host plant structured root-inhabiting AMF communities, suggesting that the plant genetic makeup may be linked to genes that control symbiosis development.

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). Following a previous Long Database Report (Dengler et al. 2018, Phyto- coenologia 48, 331–347), we provide here the first update on content and functionality of GrassPlot. The current version (GrassPlot v. 2.00) contains a total of 190,673 plots of different grain sizes across 28,171 independent plots, with 4,654 nested-plot series including at least four grain sizes. The database has improved its content as well as its functionality, including addition and harmonization of header data (land use, information on nestedness, structure and ecology) and preparation of species composition data. Currently, GrassPlot data are intensively used for broad-scale analyses of different aspects of alpha and beta diversity in grassland ecosystems.

Aim: Our aim was to quantify the extent to which nutrient pollution explains arbuscular mycorrhizal fungal community richness and composition.Location: Europe.Time period: 2014-2016.Major taxa stud ...

© 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.

Although it is well known that arbuscular mycorrhizal fungi (AMF) play a key role in the functioning of natural ecosystems, the underlying drivers determining the composition of AMF communities remain unclear. In this study, we established 138 sampling plots at 46 grassland sites, consisting of 26 acidic grasslands and 20 calcareous grasslands spread across eight European countries, to assess the relative importance of abiotic and biotic filtering in driving AMF community composition and structure in both the grassland soils and in the roots of 13 grassland plant species. Soil AMF communities differed significantly between acidic and calcareous grasslands. In root AMF communities, most variance was attributable to soil variables while very little variation was explained by host plant identity. Root AMF communities in host plant species occurring in only one grassland type closely resembled the soil AMF communities of that grassland type and the root AMF communities of other host plant species occurring in the same grassland type. The observed AMF-host plants networks were not modular but nested. Our results indicate that abiotic conditions, rather than biotic filtering through host plant specificity, are the most important drivers in shaping AMF communities in European seminatural grasslands.

Arbuscular mycorrhizal fungi (AMF) play a key role in the functioning of agricultural ecosystems. Therefore, understanding how the application of fertilizers, a common management practice, affects AMF communities is of major importance. Here we aimed to: (i) experimentally test whether different amounts and forms of phosphorus (P) fertilizer affect AMF diversity and community composition associated with the roots of apple trees (Malus domestica); (ii) identify differences in tolerance to P fertilization between AMF taxa. We used 454-pyrosequencing of the small subunit rRNA gene amplicons to quantify AMF diversity and community composition in root samples obtained from a three year field experiment, with two inorganic, three slow-release P fertilization and one control treatment. The slow-release fertilizer treatments showed significantly higher AMF richness and differed in community composition compared to the inorganic fertilizer treatments. The distribution of AMF OTUs showed a significantly nested pattern. Additionally, AMF communities in the inorganic fertilizer treatments were a subset of the communities in the slow-release fertilizer treatments. We demonstrate that application of slow-release fertilizers promoted AMF diversity in the roots of cultivated apple trees in comparison to the other treatments. The application of inorganic fertilizers elevated levels of plant-available P in the soil and selected only a small subset of abundant AMF, resulting in a lower AMF diversity. This may result in AMF communities dominated by inferior AMF mutualists.

Questions: How has vegetation species diversity and species composition changed between 1965 and 2012/13 in acidic and calcareous grasslands? What has driven this change in vegetation? Location: A 2400-km2 area around Sheffield, northern England. Methods: In 1965 a survey was conducted to describe grassland vegetation of the Sheffield region. We repeated this survey in 2012/13, revisiting acidic and calcareous grassland sites (455 quadrats). Climate, N and sulphur deposition, cattle and sheep stocking rates, soil pH, altitude, aspect and slope were considered to be potential drivers of variation in vegetation. We analysed temporal changes in vegetation and examined relationships with spatial and temporal variation in driver variables. Results: Both acidic and calcareous grasslands showed clear changes in species composition between the two time periods. In acidic grasslands there was no significant change in richness but there were declines in diversity. There were significant increases in Ellenberg N. Nitrogen deposition and grazing were identified as potential drivers of spatial and temporal patterns but it was not possible to discriminate the respective impacts of potential drivers. In calcareous grasslands there were declines in species richness, diversity and appropriate diversity indices. Climate and soil pH were identified as potential drivers of spatial and temporal patterns. Conclusions: Despite only small site losses compared to other surveys in the UK, especially within the national park, both calcareous and acidic grasslands showed very clear changes in species composition. In acidic grasslands, high abundance of Pteridium aquilinum was a particular problem and had increased considerably between the two survey periods. Atmospheric N deposition and grazing were identified as drivers of species diversity. A number of calcareous grasslands showed signs of reduced management intensity leading to scrub invasion.

Coffee is the most important tropical agricultural commodity worldwide, cultivated in more than 70 countries. Despite the plant's huge economic importance, there is very limited knowledge on the association of arbuscular mycorrhizal fungi (AMF) with coffee roots. We investigated the environmental drivers affecting the diversity and community composition of AMF on Arabica coffee in its Ethiopian center of origin. We used 454 amplicon pyrosequencing to describe AMF communities in the roots of Arabica coffee plants that were sampled along a large management intensity gradient, covering the major Ethiopian coffee production systems. We identified AMF genera that have not been reported before in Arabica coffee production regions elsewhere in the world and show that high soil phosphorus availability decreases AMF diversity and that soil pH, nitrogen and phosphorus availability strongly affect AMF community composition. At the scale of our study (max. 82 km distance between sampling points, and 770 m altitude difference), no effect was found of spatial location or altitude on AMF communities. This is the first study analyzing the drivers of naturally occurring AMF in the roots of a globally important tropical crop, providing preliminary data to improve coffee production in its native and introduced range, through targeted intervention in coffee AMF communities.

Heavy metals in urban soils may impose a threat to public health and may negatively affect urban tree viability. Vegetation spectroscopy techniques applied to bio-indicators bring new opportunities to characterize heavy metal contamination, without being constrained by laborious soil sampling and lab-based sample processing. Here we used Tilia tomentosa trees, sampled across three European cities, as bio-indicators i) to investigate the impacts of elevated concentrations of cadmium (Cd) and lead (Pb) on leaf mass per area (LMA), total chlorophyll content (Chl), chlorophyll a to b ratio (Chla:Chlb) and the maximal PSII photochemical efficiency (Fv/Fm); and ii) to evaluate the feasibility of detecting Cd and Pb contamination using leaf reflectance spectra. For the latter, we used a partial-least-squares discriminant analysis (PLS-DA) to train spectral-based models for the classification of Cd and/or Pb contamination. We show that elevated soil Pb concentrations induced a significant decrease in the LMA and Chla:Chlb, with no decrease in Chl. We did not observe pronounced reductions of Fv/Fm due to Cd and Pb contamination. Elevated Cd and Pb concentrations induced contrasting spectral changes in the red-edge (690–740 nm) region, which might be associated with the proportional changes in leaf pigments. PLS-DA models allowed for the classifications of Cd and Pb contamination, with a classification accuracy of 86% (Kappa = 0.48) and 83% (Kappa = 0.66), respectively. PLS-DA models also allowed for the detection of a collective elevation of soil Cd and Pb, with an accuracy of 66% (Kappa = 0.49). This study demonstrates the potential of using reflectance spectroscopy for biomonitoring of heavy metal contamination in urban soils. Applying leaf reflectance spectroscopy to urban trees allows for biomonitoring of heavy metal pollution and the classification of pollutants in urban soils.

Trees in urban areas face harsh environmental conditions. Ectomycorrhizal fungi (EcM) form a symbiosis with many tree species and provide a range of benefits to their host through their extraradical hyphal network. Although our understanding of the environmental drivers and large scale geographical variation of EcM communities in natural ecosystems is growing, our knowledge of EcM communities within and across urban areas is still limited. Here we characterized EcM communities using Illumina miseq sequencing on 175 root samples of the urban tree Tilia tomentosa from three European cities, namely Leuven (Belgium), Strasbourg (France) and Porto (Portugal). We found strong differences in EcM richness and community composition between cities. Soil acidity, organic matter and moisture content were significantly associated with EcM community composition. In agreement, the explained variability in EcM communities was mostly attributed to general soil characteristics, whereas very little variation was explained by city and heavy metal pollution. Overall, our results suggest that EcM communities in urban areas are significantly associated with soil characteristics, while heavy metal pollution and biogeography had little or no impact. These findings deliver new insights into EcM distribution patterns in urban areas and contribute to specific inoculation strategies to improve urban tree vitality. ispartof: FEMS Microbiol Ecol vol:94 issue:12 ispartof: location:England status: Published online

© 2018 The Authors Urban trees play a key role in mitigating environmental problems in cities, but they often face harsh environmental conditions as they generally grow in sealed soils that have small rooting space and low water availability. In this context, rapid monitoring and assessment of tree health status is critical to maintain urban trees and secure the provisioning of urban ecosystem services. Across three European cities we selected 187 Tilia tomentosa trees growing under following planting conditions: (i) sealed, trees planted in small soil pits or strips surrounded by highly sealed surfaces (concrete, pavement or asphalt); and (ii) unsealed, trees planted in roomy soil surfaces (e.g. parks). We measured leaf reflectance and fluorescence and derived a set of optical traits from the measurements. We examined whether these non-destructively measured optical traits differ between planting conditions and whether they correlate with leaf functional traits, e.g. specific leaf area (SLA), leaf water content (LWC) and leaf water per area (LWA). Compared to the unsealed trees, sealed trees showed decreased SLA and LWC while increased LWA. Leaf optical traits differed between the unsealed and sealed trees. Highly sealed soils accelerated leaf senescence of the sealed trees compared to the unsealed trees, embodied in the temporal trend of optical traits. Sealed planting conditions negatively affect urban tree health status and phenology. These negative effects can be estimated by leaf optical traits, demonstrating the great potential of optical traits in assessing tree health status. Our findings provide insights into facilitating urban green management using optical traits and remote sensing data. ispartof: ECOLOGICAL INDICATORS vol:95 pages:895-906 status: published

Questions Does addition of an arbuscular mycorrhizal fungi (AMF) inoculum increase the short-term restoration success of nutrient-poor grassland (NPG) after topsoil removal? Does distance to intact remnant grassland (IRG) patches affect restoration success, and does the effect of inoculum addition depend on distance to IRGs? Location Meerdaal forest, Oud-Heverlee, Belgium. Methods In a topsoil-removed site of 8.5 ha, where 24 IRG patches (ca. 10% of the area) were kept, 48 plots (1 m2) were established at three distances (5, 10 and 20 m) from the edge of IRG patches. Half of the plots at each distance class were inoculated with a custom-made AMF inoculum, whereas the remaining plots were used as controls. We recorded plant species abundance in the plots, just before addition of the AMF inoculum and 1 yr after. We used repeated measures ANOVA to test for effects of inoculum addition, distance to IRG patches and their interaction on plant species richness, diversity and plant community similarity between IRG patches and plots. We also evaluated the response of AMF-dependent plant species, specialist species of NPG and species with specific seed dispersal adaptations. Results Adding the inoculum positively affected species richness and/or diversity of all plant species, AMF-dependent species and specialist species. It increased plant community similarity to the IRG patches. Increasing distance from IRG patches had a negative effect on richness and/or diversity of all plant species and specialist species. The positive effect of inoculum addition on richness and/or diversity of all species, AMF-dependent species and specialist species decreased with increasing distance from IRG patches to the plots, likely indicating priority effects. Conclusions The application of a custom-made AMF inoculum increased the short-term restoration success of NPG after topsoil removal. Dispersal limitation of specialist plant species of NPG, however, likely negatively affected the effect of inoculum addition. Apart from the AMF effect, the reported strongly positive short-time effect of inoculation was likely due to the high density of IRG patches at the site, and the presence of organic matter and other micro-organisms in the inoculum.

Dissimilarity overlap curve analysis has shown that universality is a common feature in many complex microbial communities, suggesting that the same taxa interact in a similar manner when shared between communities. We present evidence that arbuscular mycorrhizal fungi, common plant root symbionts, show universal community compositions in natural ecosystems and that this pattern is conserved even at larger spatial scales. However, universality was not detected in agricultural ecosystems potentially implying that agricultural symbiont communities are formed in a different manner.

Species diversity is commonly hypothesized to result from trade-offs for different limiting resources, providing separate niches for coexisting species1-4. As soil nutrients occur in multiple chemical forms, plant differences in acquisition of the same element derived from different compounds may represent unique niche dimensions5,6. Because plant productivity of ecosystems is often limited by phosphorus7, and because plants have evolved diverse adaptations to acquire soil phosphorus6,8, a promising yet untested hypothesis is phosphorus resource partitioning6,9,10. Here, we provided two different chemical forms of phosphorus to sown grassland mesocosms to investigate phosphorus acquisition of eight plant species that are common in European grasslands, and to identify subsequent patterns of plant abundance. For the first time, we show that the relative abundance of grassland plant species can be influenced by soil phosphorus forms, as higher abundance was linked to higher acquisition of a specific form of phosphorus. These results were supported by a subsequent isotope dilution experiment using intact grassland sods that were treated with different inorganic or organic phosphorus forms. Here, 5 out of 14 species showed greater phosphorus acquisition in the inorganic phosphorus treatment, and 4 in the organic phosphorus treatments. Furthermore, for the species used in both experiments we found similar acquisition patterns. Our results support the hypothesis of phosphorus resource partitioning and may provide a new mechanistic framework to explain high plant diversity in phosphorus-poor ecosystems6,11-13. As world biodiversity hotspots are almost invariably related to phosphorus limitation8,11,12, our results may thus also be key to understanding biodiversity loss in an era of ever-increasing nutrient enrichment14.

Pollinators are declining worldwide and possible underlying causes include disease, invasive pest species and large scale land use changes resulting in habitat loss and degradation. One particular cause of habitat degradation is the increased inflow of nutrients due to anthropogenic combustion processes and large scale application of agricultural fertilizers. This nutrient pollution has been shown to affect pollinators through the loss of nectar and pollen-providing plant species. However, it may also affect pollinators through altering the nectar and pollen chemical composition of plant species, hence influencing pollinator food quality. Here, we experimentally investigated the effect of nutrient enrichment on amino acid and sugar composition of nectar and pollen in the grassland plant Sucissa pratensis, and the subsequent colony size and larval mortality of the pollinating bumblebee Bombus terrestris. We found less of the essential amino acids glycine and arginine in the pollen of fertilized plants, and more arginine, ornithine and threonine in the pollen of control plants. Nectar glucose and pollen fructose levels were lower in fertilized plants as compared to control plants. Furthermore, bumblebee colonies visiting fertilized plants showed more dead larvae than colonies visiting control plants. Our results suggest that the fitness of bumblebees can be negatively affected by changes in their food quality following nutrient pollution. If similar patterns hold for other plant and pollinator species, this may have far reaching implications for the maintenance of pollination ecosystem services, as nutrient pollution continues to rise worldwide. ispartof: PLOS ONE vol:12 issue:4 ispartof: location:United States status: published

Anthropogenic activities have severely altered fluxes of nitrogen and phosphorus in ecosystems worldwide. In grasslands, subsequent negative effects are commonly attributed to competitive exclusion of plant species following increased above-ground biomass production. However, some studies have shown that this does not fully account for nutrient enrichment effects, questioning whether lowering competition by reducing grassland productivity through mowing or herbivory can mitigate the environmental impact of nutrient pollution. Furthermore, few studies so far discriminate between nitrogen and phosphorus pollution. We performed a full factorial experiment in greenhouse mesocosms combining nitrogen and phosphorus addition with two clipping regimes designed to relax above-ground competition. Next, we studied the survival and growth of seedlings of eight common European grassland species and found that five out of eight species showed higher survival under the clipping regime with the lowest above-ground competition. Phosphorus addition negatively affected seven plant species and nitrogen addition negatively affected four plant species. Importantly, the negative effects of nutrient addition and higher above-ground competition were independent of each other for all but one species. Our results suggest that at any given level of soil nutrients, relaxation of above-ground competition allows for higher seedling survival in grasslands. At the same time, even at low levels of above-ground competition, nutrient enrichment negatively affects survival as compared to nutrient-poor conditions. Therefore, although maintaining low above-ground competition appears essential for species' recruitment, for instance through mowing or herbivory, these management efforts are likely to be insufficient and we conclude that environmental policies aimed to reduce both excess nitrogen and particularly phosphorus inputs are also necessary. ispartof: PLoS One vol:12 issue:3 ispartof: location:United States status: published

Anthropogenically increased input of nitrogen (N) and phosphorous (P) have led to a severe reduction of plant species richness in European semi-natural grasslands. Although it is well established that this species loss is not trait neutral, a thorough analysis of the effects of nutrient addition on trait based functional diversity and functional composition, independently of species loss, is lacking so far. We compiled data on the plant species abundance (releve’s) of 279 Nardus grasslands from nine European countries, across a gradient of soil N and P content. Functional diversity (Petchy and Gaston’s FDc, weighted FDc and quadratic entropy) and mean trait composition were calculated for each releve, based on 21 functional traits. Differences in functional diversity and functional composition were related to differences in soil N, atmospheric N deposition, soil P and pH, while controlling for geographic location and species richness. All functional diversity measures decreased with increasing soil N, with wFDc also decreased by soil P, independent of species loss. This was accompanied by clear shifts in functional trait composition, associated with shifts from below-ground competition for nutrients to above-ground competition for light. This resulted in a decrease in insect-pollinated therophytes and chamaephytes and an increase in long-lived, clonal graminoids and hemicryptophytes under increasing soil N and P. These functional community changes can be expected to alter both ecosystem functioning and service provisioning of the studied grasslands. Our research emphasizes the importance of a reduction of both N and P emission throughout Europe for sustainable conservation of these communities.

Aim Although many studies support the prevailing paradigm of nitrogen (N)-driven biodiversity loss, some have argued that phosphorus (P) may be the main culprit. This questions the generality of the global threat through N enrichment. The major objective here was to quantify the relative importance of soil N and P in explaining patterns of plant species richness, under different levels of N and P limitation. Location North-western Europe. Methods We collected soil, productivity and plant species data from 132 semi-natural grasslands located along a gradient of nutrient availability and atmospheric N deposition. We used linear mixed models to investigate the relation between soil nutrients, acidity, limitation and productivity on one side, and indices for plant species richness on the other. Results Mixed models explained between 38 and 50% of the total variation in species numbers, forbs and endangered species. Soil P was significantly negatively related to total species number, forbs and endangered species. Soil N was only significantly negatively related to number of forbs and endangered species. Compared with soil P, the explained variation attributed to soil N was between five- and twenty-fold lower. P-limited grasslands exhibited higher species richness, numbers of forbs and endangered species. Species richness and number of forbs decreased with lower soil acidity. N deposition was negatively related to the number of forbs and endangered species, as well as to soil acidity. Productivity was weakly positively related to soil P and negatively to species and forb numbers. We found no interaction factors between the explanatory variables. Main conclusions P enrichment can present a greater threat to biodiversity than N enrichment in at least some terrestrial ecosystems. However, as N- and P-driven species loss appeared independent, our results suggest that simultaneously reducing N and P inputs is a prerequisite for maintaining maximum plant diversity.

Summary 1. Both nitrogen (N) and phosphorus (P) enrichment have been identified to drive plant species losses from nutrient-poor semi-natural grasslands. The relative contribution of N vs. P to species loss remains unclear, however. 2. We investigated how soil N and P availability affect the occurrence of 61 grassland species across North-western Europe. We selected 132 study sites, located in Great Britain, Belgium and France, along a soil fertility gradient based on variability in atmospheric N deposition and on nutrient input from adjacent agricultural land. To gain insight into the underlying ecological mechanisms of species loss, we examined the role of a suite of plant traits that may mediate a species’ response to increased N or P availability. 3. Mixed logistic regression showed that the occurrence of 24 plant species (39AE3%) was affected by soil nutrient availability. Of these species, 18 were negatively affected by increased P (29AE5%) and five by increased N (8AE2%). Regionally declining plant species were absent from both P-rich and N-rich grasslands. Higher susceptibility to elevated P was associated with stress tolerance, low maximum canopy height and symbiosis with arbuscular mycorrhizae. 4. Synthesis and applications. Although we also identified negative effects on plant diversity through N enrichment, our results strongly suggest that P enrichment is a more important driver of species loss from semi-natural grasslands. Species in symbiosis with mycorrhizae and with low canopy height are especially at risk. Because detrimental effects of P enrichment are very difficult to mitigate due to the persistence of P in the soil, nature management should give absolute priority to preventing P input in grasslands through fertilization, agricultural run-off or inundation with P-polluted surface water. To restore species-rich grasslands on P-enriched soils, top soil removal appears crucial and more research regarding alternative removal strategies is essential.

Floral traits have evolved to maximize reproductive success by attracting pollinators and facilitating pollination. Highly attractive floral traits may, however, also increase the degree of self-pollination, which could become detrimental for plant fitness through inbreeding depression. Floral nectar is a trait that is known to strongly mediate pollinator attraction and plant reproductive success, but the particular role of the nectar amino acid (AA) composition is poorly understood. Therefore, we experimentally manipulated the nectar AA composition and abundance of the Lepidoptera-pollinated orchid Gymnadenia conopsea through soil fertilization, and we quantified AA content and AA composition through high performance anion exchange chromatography with pulsed amperometric detection. Mixed models were then used to evaluate differences in pollinia removal, fruit set, seed set and degree of selfing between fertilized and control individuals. Selfing rates were estimated using microsatellite markers. We found that fertilized individuals had a significantly higher nectar AA content and an altered AA composition, whereas plant height, number of flowers, nectar volume and sugar concentration remained unchanged. Fertilized individuals also had significantly more pollinia removed and a higher fruit set, whereas control plants that did not receive the fertilization treatment had significantly fewer selfed seeds, and more viable seeds. Although we cannot exclude a role of changes in floral scent following the fertilization treatment, our results strongly suggest a relation among nectar AA composition, fruiting success and selfing rates. Our results also indicate potential consequences of nutrient pollution for plant reproductive success, through the induced changes in nectar AA composition.

Species-rich grassland communities are one of the most important habitats for biodiversity and of high conservation priority in Europe. Restoration actions are mainly focused on the improvement of abiotic conditions, such as nutrient depletion techniques, and are generally based on the assumption that the target community will re-establish at the restored site when the target species exist in the neighborhood. Information on the contemporary seed-dispersal range is therefore crucial to develop effective conservation measures. Here, we investigated the contemporary long-distance seed dispersal and genetic structure of the grassland herb Thymus pulegioides in an intensively managed agricultural landscape in Flanders (Northern Belgium). Assignment tests based on amplified fragment length polymorphisms revealed very low levels of effective seed dispersal between populations although seed availability and seed viability was not a limiting factor. The process of fragmentation has resulted in a high population differentiation and without further incoming gene flow the remnant populations are prone to further genetic erosion and perhaps extinction. Our findings illustrate that restoring suitable abiotic habitat conditions in the neighborhood of existing populations does likely not guarantee colonization for this grassland specialist. For the survival of the species, existing populations should be functionally connected and seed addition may be necessary for successful conservation to overcome dispersal-limitation. ispartof: Conservation Genetics vol:16 issue:4 pages:765-776 status: published

International audience; Nutrient pollution presents a serious threat to biodiversity conservation. In terrestrial ecosystems, the deleterious effects of nitrogen pollution are increasingly understood and several mitigating environmental policies have been developed. Compared to nitrogen, the effects of increased phosphorus have received far less attention, although some studies have indicated that phosphorus pollution may be detrimental for biodiversity as well. On the basis of a dataset covering 501 grassland plots throughout Europe, we demonstrate that, independent of the level of atmospheric nitrogen deposition and soil acidity, plant species richness was consistently negatively related to soil phosphorus. We also identified thresholds in soil phosphorus above which biodiversity appears to remain at a constant low level. Our results indicate that nutrient management policies biased toward reducing nitrogen pollution will fail to preserve biodiversity. As soil phosphorus is known to be extremely persistent and we found no evidence for a critical threshold below which no environmental harm is expected, we suggest that agro-environmental schemes should include grasslands that are permanently free from phosphorus fertilization.