Decreasing nitrogen deposition rates: Good news for oligotrophic grassland species?

Several studies have found that increased nitrogen (N) deposition leads to a decline in species richness in semi-natural grasslands, mainly due to the loss of species typical of nutrient-poor soils. However, after reaching a peak around 1990, N deposition has decreased in Europe over the last 30 years. In this study, we investigated the changes in species number and composition of semi-natural grasslands during this period of declining N deposition. To this end, we compared the data from the first survey (2001-2005) of 147 grassland sites in Switzerland with those from the third survey (2011-2015). We further analysed the vegetation development of a specific hay meadow from 1992 to 2013. In this grassland, total vegetation cover and the cover of graminoid species decreased, while the cover of oligotrophic species increased. At the 147 grassland sites, total species number decreased at sites with still high levels of N deposition and it tended to increase at sites with low N deposition, i. e. below the critical load for N deposition. The number of oligotrophic grassland species increased at sites with a large decrease in N deposition and strong inclination. Thus, the results of this study indicate that the reduction of N emissions had a measurable positive effect on species diversity in these semi-natural grasslands. Most of the grasslands surveyed appear to be quite resilient against N deposition, i. e. they do not shift to an alternative low diversity state dominated by a few competitive species, and recovery of the species composition as a result of the decrease in N deposition seems possible, especially on steep slopes. Furthermore, the study underlines the importance of regular management of semi-natural, unfertilised, low-productivity grassland to maintain the diversity of oligotrophic grassland species.     

Long-term species change in a Uruguayan grassland

Abstract. This paper reports on species changes which occurred in plots in a Uruguayan grassland after 55 yr of continuous grazing. The study was based on the pioneering work of B. Rosengurtt who sampled 10 plots in 1935; in 1990 we re-sampled five of them. Comparative studies were based on rank-frequency, diversity, evenness and multivariate analyses. Diversity was measured as species richness S, Shannon's index H and its exponential form N = e^H, evenness E = N/S and Gini's coefficient of hierarchy G. Species-richness of the plots increased significantly. Species which entered the sites are mostly weedy species, while those that have decreased are mostly palatable forage species. Evenness, however, remained constant. Floristic change was chiefly associated with time, and secondarily with plot soil-type. During the 55 yr that elapsed between both observation periods, a dramatic floristic change has occurred in this grassland site.     

Long-term after effects of fertilization on above-ground phytomass and species diversity in calcareous grassland

Abstract. Long-term after effects on species number and productivity in calcareous grassland were analyzed after cessation of fertilization. Three series of permanent plots were monitored yearly from 1971 to 1993. These series differed in duration of fertilization and in fertilizer composition, notably the amount of nitrogen and phosphorus. Yearly above-ground production decreased in all series after fertilization had been stopped, however at different speeds. The grass/forb ratio also decreased, while species number and Shannon index of diversity (H') increased. In the series where fertilizer treatment was stopped in 1967 (1968-series), productivity and grass/forb ratio decreased equally as in the plots where fertilization with a high nitrogen content was stopped in 1979 (80/Npk-series). However, species number in the plots of the 80/Npk-series increased faster than in the 1968-series. This was probably the result of a higher number of species present in the seed rain from the surrounding vegetation in the period after 1980 than from after 1968. In the plots receiving a high amount of phosphorus (80/nPk series), the productivity decreased more slowly than in the high nitrogen plots (80/Npk-series). At approximately the same total production level, the grass/forb ratio remained higher for another five-year period in the 80/nPk-series. Species number and the Shannon index of diversity increased more slowly in the 80/nPk-series than in the 80/Npk-series. Over a given range of productivity, changes in species number and Shannon index correlated better with the grass/forb ratio than with total above-ground phytomass. Therefore, restoration of species-rich grassland should not only be focused on lowering the yearly production, but also on reducing the grass component of the vegetation.     

Nitrogen deposition causes widespread loss of species richness in British habitats

We use national scale data to test the hypothesis that nitrogen (N) deposition is strongly negatively correlated with plant species richness in a wide range of ecosystem types. Vegetation plots from a national ecological surveillance programme were drawn from heathland, acid, calcareous and mesotrophic grassland habitats. Mean species number and mean plant traits were calculated for each plot and related to atmospheric N deposition. There was a significant reduction in species richness with N deposition in acid grassland and heathland even after fitting covarying factors. In acid grassland and heathland, evidence from trait changes suggested that acidification rather than increased fertility was responsible for species loss. In contrast, calcareous grassland showed evidence of eutrophication in response to increasing N deposition. Loss of species richness from chronic N deposition is apparent in infertile grasslands and heathland. Mechanisms associated with loss of species richness differ between habitats so mitigation of N deposition should be targeted to habitat type.     

Missing the target? A critical view on butterfly conservation efforts on calcareous grasslands in south-western Germany

Butterflies are strongly declining on grassland habitats of Central Europe. Therefore, the success of conservation measures on high quality grassland habitats is controversially discussed. We compared the changes in butterfly diversity and community structure on six managed calcareous grasslands with eight unmanaged vineyard fallows. We obtained strong losses of species diversity and remarkable shifts of community compositions on both habitat types. However, the changes on vineyard fallows were only slightly more severe but more stochastic than on the calcareous grasslands. The shifts in community composition with respect to functional species traits were rather similar between the two different grassland types so that complex butterfly communities evolved into generalist-dominated ones. Connectivity was higher among vineyard fallows than among calcareous grasslands. Consequently, conservation measures on calcareous grasslands only partly achieved their goal to maintain the high species diversity and functional complexity still observed in the 1970s. The negative impacts of eutrophication and monotonisation of the landscape as well as climate change are affecting all habitats, independently from management concepts. Therefore, management on conservation sites can buffer against these effects, but is not sufficient for a full compensation.     

中国东北样带草地群落放牧干扰植物多样性的变化

放牧干扰是草地群落植物多样性变化的主要影响因素之一。中国东北样带9个草地群落放牧干扰植物多样性变化的研究结果表明：中牧或重牧阶段Shannon指数达最大值,形成中牧（重牧）>重牧（中牧）>轻牧>过牧的规律。群落物种丰富度、均匀度与多样性的相关分析表明,均匀度变化对多样性变化具有更大的贡献率,而丰富度呈下降趋势,即轻牧（中牧）>中牧（轻牧）>重牧>过牧。生活型功能群多样性也表现出明显的变化。中国东北样带草地群落植物多样性的分布格局是：草甸草原>典型草原>典型草原 >荒漠草原>碱化草原,并且群落物种丰富度对多样性有更大贡献率。     

中国东北样带草地群落放牧干扰植物多样的变化

放牧干扰是草地群落植物多样性变化的主要影响因素之一。中国东北样带９个草地群落放牧干扰植物多样性性变化的研究结果表明：中牧或重牧阶段Ｓhannon指数达最大值。形成中牧（重牧）＞重牧（中牧）＞轻牧＞过牧的规律。群落物种丰富度、均匀度与多样性的相关分析表明,均匀度变化对多样性变化具有更大的贡献率。而丰富度呈下降趋势,即轻牧（中牧）＞中牧（轻牧）＞重牧＞过牧。生活型功能群多样性也表现出明显的变化。中国东北样带草地群落植物多样性的分布格局是：草甸草原＞典型草原＞典型草原＞荒漠草原＞碱化草甸,并且群落物种丰富度对多样性有更大贡献率。     

半干旱黄土区不同管理措施下草地群落结构对短期氮、水添加的响应

氮沉降和降水变异显著影响草地群落结构和功能,但缺乏对不同管理措施下草地群落结构对氮沉降和降水变异响应的研究。为模拟不同管理措施下草地群落结构对氮沉降和降水变异的响应特征,以半干旱黄土区云雾山国家自然保护区典型草原为研究对象,系统分析了在封育、刈割和火烧三种管理措施下,氮添加和水添加对群落地上生物量、功能群组成和群落多样性的影响。结果表明,氮添加和水添加对地上生物量、功能群组成和群落多样性指数的影响因管理措施不同有所差异。（1）在封育草地上,氮添加显著降低物种多样性,对地上生物量影响较小;水添加显著增加物种多样性指数,氮添加和水添加的交互作用显著增加地上生物量、禾本科所占比例和莎草科所占比例;物种多样性指数均与地上生物量无显著相关,与不同功能群所占比例显著相关。（2）在刈割草地上,氮添加和水添加显著提高草地群落地上生物量,氮添加和水添加交互作用尤为显著;氮添加和水添加显著增加物种丰富度指数,对物种均匀度影响较小;杂草类所占比例和地上生物量对Shannon-Weiner多样性指数的贡献率较大。（3）在火烧草地上,氮添加和水添加显著提高群落地上生物量,对物种多样性的影响因年份不同有所差异,氮添加和水添加交互作用具有累加效应;Shannon-Weiner多样性指数与地上生物量呈显著负相关,与莎草科所占比例呈显著正相关。研究表明管理措施显著影响群落结构对氮添加和水添加的响应特征,亦改变生产力和物种多样性的关系模式,为更好地应对全球变化进行草地管理提供数据支撑。     

Diversity patterns in sandy forest-steppes: a comparative study from the western and central Palaearctic

The Palearctic forest-steppe biome is a narrow vegetation zone between the temperate forest and steppe biomes, which provides important habitats for many endangered species and represents an important hotspot of biodiversity. Although the number of studies on forest–grassland mosaics is increasing, information currently available about the general compositional and structural patterns of Eurasian forest-steppes is scarce. Our study aimed to compare the habitat structure, species composition and diversity patterns of two distant sandy forest-steppes of Eurasia. We compared 72 relevés made in the main habitat components (forest, forest edge and grassland) of sandy forest-steppes in three Hungarian and three Kazakh sites. The size of the plots was 25 m². Species number, Shannon diversity and species evenness values were calculated for each plot. Fidelity calculations and linear mixed effects models were used for the analyses. We found that the vegetation and diversity patterns of the two forest-steppes are similar and their components play important roles in maintaining landscape-scale diversity. Despite the higher species richness in Hungary, Shannon diversity was higher in Kazakhstan. The deciduous forest edges of both areas had significantly higher species richness than the neighbouring habitats (forests and grasslands); therefore they can be considered local biodiversity hotspots. Due to the special characteristics of this vegetation complex, we emphasize the high conservation value of all landscape components as a coherent system throughout the entire range of the Eurasian forest-steppe biome.     

Is the relation between nutrient supply and biodiversity co‐determined by the type of nutrient limitation?

Correlative studies have shown a ‘hump‐backed’ relation between the vegetation N:P ratio and plant species diversity with the highest diversity at balanced N:P ratios (between 10 and 14). We tested the hypothesis that adding growth‐limiting nutrients to mesotrophic grasslands that were in shortage of either N (N:P ratio<10) or P (N:P ratio>14) would lead to an increase of plant diversity. Thereto, we studied the effects of long‐term (11 yr) experimentally increased N and/or P supply on soil nutrient pools, vegetation nutrient dynamics and biodiversity in a riverine grassland in the Netherlands with a low soil N:P ratio (N shortage) and a peat grassland with a high soil N:P ratio (P shortage), respectively. Eleven years of nutrient addition hardly had any effects on the total stocks of C, N and P in the soils of both sites, due to the large size of the soil nutrient pools already present and to the management at both sites (annual hay‐making and ‐removal). However, in the riverine grassland the treatments increased the cycling of the small pool of labile N and P compounds resulting in large increases in annual fluxes of especially N. In the unfertilised controls, species establishments balanced more or less species losses during an 11 year period, thus leading to a dynamic equilibrium of the species pool. However, contrary to our hypothesis, addition of the growth‐limiting nutrient led at both sites to a reduction of species diversity even when total biomass remained below critical levels. Species diversity and species evenness were strongly determined by N mineralisation and to a lesser extent by total soil N and extractable P, respectively. Total aboveground biomass of the vegetation was determined by total soil N. Our study shows that patterns found in correlative studies of the relation between plant diversity and soil and vegetation N:P ratio can not be translated into successful experimental manipulations to enhance biodiversity. The most likely explanation is that colonization limitation occurred in the fertilized plots and that not sufficient diaspores of potentially new species could reach and/or colonize the plots to compensate for the species extinctions as a result of increased nutrient supply.     

The effects of quantity and duration of simulated pollutant nitrogen deposition on root-surface phosphatase activities in calcareous and acid grasslands: a bioassay approach

A field and laboratory based bioassay has been developed to investigate the effects of the quantity and duration of simulated pollutant nitrogen (N) deposition on root-surface phosphomonoesterase (PME) activities in calcareous and acid grasslands. Seedlings of Plantago lanceolata were transplanted to a calcareous grassland and Agrostis capillaris seedlings were grown in microcosms containing soil from an acid grassland that had received either 7 yr (long-term) N additions or 18 months (short-term) N and phosphorus (P) additions. The bioassay revealed that short-term N treatments had little effect on the enzyme activity, whereas long-term N additions significantly increased PME activity within 7 d of transplanting into the field plots. Root-surface PME activity of A. capillaris was significantly reduced in soil that received additions of P. In the plots receiving long-term additions of N, a strong relationship was observed between extractable soil ammonium and root-surface PME activity. Soil ammonium concentrations accounted for 67% of the variation in PME activity of P. lanceolata in the calcareous grassland, and 86% of the variation in PME activity of A. capillaris in the acid grassland. These results provide evidence that N deposition may have considerable effects on the demand and turnover of P in ecosystems that are approaching or have reached N saturation.     

陇东黄土高原丘陵沟壑区天然草地植物多样性研究

以陇东黄土高原丘陵沟壑区天然草地30个植被样点的样方资料为依据,以相对生物量为指标,对该地区群落植物多样性进行了研究。结果表明,该地区天然草地可划分为长芒草草原、次生杂类草草原和杂类草草甸草原3个植被类型,16个群落按放牧程度分为禁牧草地、轻牧草地和过牧草地3个层次。多样性指数显示草甸草原>长芒草草原>次生杂类草草原,轻牧草地>禁牧草地>过牧草地的趋势。放牧程度可能是影响该区域天然草地植物多样性的关键因素,出现海拔增高和降水增加多样性指数降低的趋势可能是放牧干扰的结果。相关性分析显示物种的Shannon-wiener和Simpson多样性指数与Shannon和Simpson均匀度指数间均呈显著正相关,而与生物量之间呈显著负相关。     

Hybrid ecosystems can contribute to local biodiversity conservation

Calcareous grasslands have become severely threatened habitats in Europe. The aim of this study was to investigate the changes in plant species richness, and functional and phylogenetic diversity in northern Estonian calcareous (alvar) grasslands resampled after 90 years of land-use change. Functional traits characterizing species that have benefited most from decreased habitat area and altered environmental conditions, and additional species that can potentially inhabit the remaining grassland patches were identified. Also changes in the relative amount of habitat-specific species were studied to detect a possible decrease in habitat integrity. Although grasslands in the studied region had lost most of their original area (~90 %), species richness had substantially increased due to invasion by more competitive, nutrient-demanding native species. Functional diversity generally increased, whereas phylogenetic diversity showed no response to altered conditions. Overall, these grasslands have lost their integrity as calcareous grassland habitat type in the region, because the relative amount of habitat-specific characteristic species has declined significantly. However, although the grasslands have transformed to a ‘hybrid’ habitat type and restoration to their previous state is likely not reasonable, such degraded species-rich grassland fragments can still be recognized as important habitats to preserve high local biodiversity and several characteristic species of calcareous grasslands. As current landscapes consist of an increasing number of hybrid and novel communities, new tools to supplement traditional conservation or restoration practices are necessary to recognize and maintain regions and habitats of high local biodiversity.     

Small mammal herbivores mediate the effects of soil nitrogen and invertebrate herbivores on grassland diversity

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Vegetation monitoring in a 100-year-old calcareous grassland reserve in Germany

Calcareous grasslands harbor specialized species and are cultural relics. Therefore, they are prime habitates for conservation and restoration, but negative effects of inappropriate management, eutrophication and fragmentation continue to exist. These effects also influence grasslands which serve as target for restoration. Unfortunately, monitoring of long-term vegetation dynamics in calcareous grassland is rare. Here, we studied such changes over 35 years in the nature reserve ‘Garchinger Heide’, which is well known for its high abundance of rare species. Furthermore, it has been managed for conservation for more than 100 years. Therefore, species composition, total species richness, numbers of habitat specialists, red-list species and the proportion of graminoids were examined in 42 plots with frequency recording (1984–2018), and in 40 plots based on vegetation relevés (2003–2018). Ellenberg indicator values, specific leaf area, seed mass, and canopy height were analysed to detect patterns in trait response to environmental change. Within 35 years there were considerable vegetation dynamics. Specialist plants of calcareous grassland and red list species decreased, and insect-pollinated species declined in contrast to wind-pollinated species. Ellenberg N as well as graminoid abundance, canopy height, seed mass, and multi-trait functional dispersion increased, while specific leaf area showed no such change. Our results suggest that environmental change like deposition of atmospheric nitrogen, management regime, pollinator decline or isolation could be correlated with vegetation dynamics, while these correlations would need experimental confirmation. The grassland management certainly helped achieving several conservation goals, although it was not able to stop a decrease of rare species. The results show that also in nature reserves with long-term conservation management monitoring is essential to detect vegetation dynamics and to adjust the management to these changes.     

扎龙湿地不同生境的昆虫多样性

为了探讨湿地不同生境对昆虫物种多样性的影响,对扎龙湿地8种生境的昆虫进行了系统调查.共捕获昆虫5822只,分属11目58科143种,其中直翅目、双翅目、蜻蜒目为扎龙湿地的优势类群.不同生境中,草原草甸昆虫多样性最高,湖边生境多样性指数和均匀度指数均较高,杂草甸均最低.聚类分析和主分量分析结果表明,不同生境的昆虫群落相似性与水资源状况和植被类型有关,捕食性类群种类数和个体数量对昆虫群落稳定性具有重要的调控作用.湖边生境昆虫群落稳定性最强,湿草甸稳定性最弱.湿地水资源状况能影响昆虫生存生境,进而影响昆虫群落的组成和分布格局.     

Long‐term after‐effects of fertilisation on the restoration of calcareous grasslands

Question: What are the long‐term implications of former fertilisation for the ecological restoration of calcareous grasslands? Location: Gerendal, Limburg, The Netherlands. Methods: In 1970, ten permanent plots were established in just abandoned agricultural calcareous grassland under a regime of annual mowing in August. From 1971 to 1979, two different fertiliser treatments were applied twice a year to a subset of the plots (artificial fertiliser with different proportions of nitrogen and phosphorus). The vegetation of the plots was recorded yearly and vegetation biomass samples were taken for peak standing crop and total amounts of nitrogen, phosphorus and potassium. Species composition and floristic diversity were analysed over the research period (1970–2006) and between the treatments, including the use of multivariate techniques (Detrended Correspondence Analysis). Results: In terms of species number, there is a clear optimum 10 to 20 years after fertilisation has been terminated. Afterwards, there is a slow decrease; no new species appear and species of more nutrient‐rich conditions gradually disappear. For the fertilised plots that received a relatively high proportion of N, effects are found only in the first years, whereas, for the plots that received a relatively high proportion of P, long‐term after‐effects are found in species composition, peak standing crop, total amounts of phosphorus in biomass, and in soil phosphorus data. Conclusions: The effect of artificial fertiliser with a large amount of nitrogen disappears in less than ten years when mown in August, including removal of the hay. This is a promising result for restoration of N‐enriched calcareous grasslands, as the applied dose of nitrogen in this experiment largely exceeds the extra input of nitrogen via atmospheric deposition. Application of fertiliser with a large amount of phosphorus, however, has effects even more than 25 years after the last addition. There are no prospects that this effect will become reduced in the near future under the current mowing management.     

Impacts of ecosystem engineers on community attributes: effects of cushion plants at different elevations of the Chilean Andes

Ecosystem engineers are organisms able to modulate environmental forces and, hence, may change the habitat conditions for other species. In so doing, ecosystem engineers may affect both species richness and evenness of communities and, in consequence, change species diversity. If these changes in community attributes are related to the magnitude of the habitat changes induced by the engineers, it seems likely that engineer species will have greater effects on diversity in sites where they cause larger habitat changes. We addressed this issue by evaluating the effects of three alpine cushion plants on species richness, evenness, and diversity of high-Andean plant communities. Given that the difference in microclimatic conditions between cushions and the external environment increases with elevation, we proposed that these organisms should have greater effects on community attributes at higher than at lower elevation sites. Results showed that the three cushion species had positive effects on species richness, diversity, and evenness of plant communities. It was also observed that the magnitude of these effects changed with elevation: positive effects on species richness and diversity increased towards upper sites for the three cushions species, whereas positive effects on evenness increased with elevation for one cushion species but decreased with elevation for other two cushion species. These results suggest that the presence of cushions is important to maintain plant diversity in high-Andean communities, but this positive effect on diversity seems to increase as the difference in environmental conditions between cushions and the external environment increases with elevation.     

Langfristige Auswirkungen ehemaliger Bewirtschaftungsvarianten auf die Diversit?t von Blütenpflanzen, Moosen und Flechten eines Kalkmagerrasens

Jeschke M., Kiehl K., Pfadenhauer J. and Gigon A. 2008. Long-term effects of former management on the diversity of vascular plants, mosses and lichens in a calcareous grassland. Bot. Helv. 118: 95 – 109. Long-term effects of grassland management on the species richness of vascular plants, mosses and lichens were studied in a calcareous grassland near Schaffhausen (Switzerland). Experimental plots were treated for 22 years (1977 – 1999) with different mowing regimes, annual burning in late winter, or abandonment. From 2000 onwards, all plots were mown annually in autumn. Five years later (2004), we studied the after-effects of the previous treatments by recording the cover of vascular plants, mosses and lichens in nested plots with sizes from 0.01 m² to 16 m². Vascular plant species richness as well as the number of species characteristic for calcareous grasslands were significantly lower in formerly abandoned or burnt plots than in regularly mown plots, with larger differences on small subplots. There was no significant effect of the previous mowing time (July vs. October) or mowing frequency (annual or every two years) on vascular plants. Mosses and lichens had been eliminated by annual burning between 1977 and 1999, and only few moss species re-colonized the formerly burnt plots until 2004. The species richness of mosses did not differ between formerly abandoned and mown plots, but there were more mesophytic and fewer xerophytic species in the formerly abandoned plots. Mesophytic mosses were also more abundant in plots that used to be mown every second year than in those mown annually. Our data indicate that even after five years of similar management, i.e. annual mowing in autumn, the previous long-term management had strong after-effects on species richness and species composition of calcareous grasslands. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. Eingereicht am 19. M?rz 2008, Angenommen am 30. August 2008, Redaktion: Sabine Güsewell     

Soil Disturbance as a Grassland Restoration Measure—Effects on Plant Species Composition and Plant Functional Traits

Soil disturbance is recognized as an important driver of biodiversity in dry grasslands, and can therefore be implemented as a restoration measure. However, because community re-assembly following disturbance includes stochastic processes, a focus only on species richness or establishment success of particular species will not inform on how plant communities respond ecologically to disturbance. We therefore evaluated vegetation development following disturbance by quantifying species richness, species composition and functional trait composition. Degraded calcareous sandy grassland was subjected to experimental disturbance treatments (ploughing or rotavation), and the vegetation was surveyed during four subsequent years of succession. Treated plots were compared with control plots representing untreated grassland, as well as nearby plots characterized by plant communities representing the restoration target.Species richness and functional diversity both increased in response to soil disturbance, and rotavation, but not ploughing, had a persistent positive effect on the occurrence of specialist species of calcareous sandy grassland. However, no type of soil disturbance caused the plant species composition to develop towards the target vegetation. The disturbance had an immediate and large impact on the vegetation, but the vegetation developed rapidly back towards the control sites. Plant functional composition analysis indicated that the treatments created habitats different both from control sites and target sites. Community-weighted mean Ellenberg indicator values suggested that the observed plant community response was at least partially due to an increase in nitrogen and water availability following disturbance. This study shows that a mild type of disturbance, such as rotavation, may be most successful in promoting specialist species in calcareous sandy grassland, but that further treatments are needed to reduce nutrient availability. We conclude that a functional trait based analysis provides additional information of the vegetation response and the abiotic conditions created, complementing the information from the species composition.