Inconsistent detection of extinction debts using different methods

The extinction debt, delayed species extinctions following landscape degradation, is a widely discussed concept. But a consensus about the prevalence of extinctions debts is hindered by a multiplicity of methods and a lack of comparisons among habitats. We applied three contrasting species–area relationship methods to test for plant community extinction debts in three habitats which had different degradation histories over the last century: calcareous grassland, heathland and woodland. These methods differ in their data requirements, with the first two using information on past and current habitat area alongside current species richness, whilst the last method also requires data on past species richness. The most data‐intensive, and hence arguably most reliable method, identified extinction debts across all habitats for specialist species, whilst the other methods did not. All methods detected an extinction debt in calcareous grassland, which had undergone the most severe degradation. We conclude that some methods failed to detect an extinction debt, particularly in habitats that have undergone moderate degradation. Data on past species numbers are required for the most reliable method; as such data are rare, extinction debts may be under‐reported.     

Ecological effects of atmospheric reactive nitrogen deposition on semi-natural terrestrial ecosystems

Evidence that enhanced reactive nitrogen deposition is affecting semi-natural terrestrial ecosystems comes from historic increases in plant tissue N concentrations, correlations between tissue N concentrations and present-day total atmospheric N deposition, changes in plant amino-acid composition and effects on N assimilation. The ecological significance of such changes in biomarkers is uncertain. This paper explores the ecological significance of reactive atmospheric N deposition through a review of previous experimental findings and new experimental evidence from an acidic and a calcareous grassland, both showing phosphorus limitation, and a N-limited Calluna vulgaris (L.) Hull heathland in upland Britain. Nitrogen addition in the range 0–20 g N m⁻² yr⁻¹ initially (years 0–4) increased the growth of Calluna and a decline in some subordinate species. In subsequent years, shoot extension was not stimulated, but winter injury was observed from 1993 onwards, suggesting a strong interaction between N supply and climatic conditions. By contrast, the grasslands showed a small decrease in the cover of higher plants in later years (6–7) of the experimental treatments (0–14 g N m⁻² yr⁻¹) and no growth stimulation. All N treatments reduced the bryophyte cover in the acidic grassland. There were marked effects on below-ground processes, including a sustained stimulation of N mineralization in the grassland soils, and an increase in the bacterial utilization of organic substrates in the heathland, as measured in BIOLOG plates. The results strongly suggest the importance of atmospheric N deposition on microbially driven processes in soils, and are discussed in relation to the scale of potential ecosystem changes and their reversibility by pollution abatement.     

Heat shock increases the reliability of a temperate calcareous grassland seed bank study

Reliable assessment of the density and species richness of the viable seeds in the soil is essential to estimate the probability of successful restoration of a particular plant community. Since temperate calcareous grasslands contain several thermophilous species typical of fire prone habitats, heat shock can be expected to break dormancy of target species for calcareous grassland restoration. We tested the effect of heat shock on the estimated density and species richness of seeds in soil samples. Heat shock treatments between 70 and 110°C resulted in both a higher estimated seed density and estimated species richness, in particular of specialist calcareous grassland species. Moreover, 25 species germinated exclusively from the heat-treated samples. This indicates that several species would have been missed by handling soil seed bank samples following the standard germination protocol, resulting in an underestimation of the restoration potential.     

Soil pH and phosphorus drive species composition and richness in semi-natural heathlands and grasslands unaffected by twentieth-century agricultural intensification

Background: Increased soil phosphorus (P) caused by agricultural intensification has been associated with decreased plant species richness (SR) in central Europe. How plant communities and soil P gradients are related in unimproved open habitats remains unclear.

Aims: The aim of this article was to characterise the relationship between soil chemical parameters and plant species composition and richness in unimproved open habitats.

Methods: The influence of soil chemical parameters (pH, P, K, Mg) on species composition was assessed, using data from 40 heathland and 54 grassland plots, by non-metric multidimensional scaling and permutational multivariate analysis of variance. The relationship between soil chemical parameters and SR was tested by linear mixed effects models.

Results: A direct relationship between heathland community composition and pH was observed, explaining 10% of variation in species composition, while P, Mg and pH together explained 17% of variation in grassland composition. In heathlands, SR increased with increasing pH, whereas in grasslands, SR decreased with increasing soil P.

Conclusions: Soil chemical parameters were substantially related to plant community composition and richness. In an area spared from a century of agricultural intensification, reduced pH appeared to constrain SR in heathlands, while even slight P increases (<10 mg kg^?1) depressed plant SR in semi-natural grasslands.     

Direct and indirect effects of nitrogen deposition on species composition change in calcareous grasslands

Atmospheric nitrogen (N) deposition has been identified as a major threat to biodiversity, but field surveys of its effects have rarely focussed on sites which are actively managed to maintain characteristic species. We analysed permanent quadrat data from 106 plots in nature reserves on calcareous grassland sites in the United Kingdom collected during a survey between 1990 and 1993 and compared the data with the results from resurvey of 48 of these plots between 2006 and 2009. N deposition showed no significant spatial association with species richness, species diversity, or the frequency of species adapted to low nutrient conditions in the 1990–1993 dataset. However, temporal analysis showed that N deposition was significantly associated with changes in Shannon diversity and evenness. In plots with high rates of N deposition, there was a decrease in species diversity and evenness, a decline in the frequency of characteristic calcareous grassland species, and a lower number of rare and scarce species. As all sites had active management to maintain a high diversity and characteristic species, our results imply that even focussed management on nature conservation objectives cannot prevent adverse effects of high rates of N deposition. Structural equation modelling was used to compare different causal mechanisms to explain the observed effects. For change in Shannon diversity, direct effects of N deposition were the dominant mechanism and there was an independent effect of change in grazing intensity. In contrast, for change in herb species number, indirect effects on soil acidity, linked to both N and S deposition, were more important than direct effects of N deposition.     

Habitat associations of epigeal spiders in upland calcareous grassland landscapes: the importance for conservation

Upland calcareous grassland landscapes are typically comprised of a matrix of calcareous grassland, acid grassland and limestone heath plant communities. This matrix of habitats is produced by a combination of underlying geology, climate and management. These landscapes are typically managed through grazing, with management targeted to maintain particular plant communities in the calcareous grassland habitat, whilst patches of acid grassland and limestone heath are not targeted by conservation management. The biodiversity value of acid grassland and limestone heath patches within the calcareous grassland matrix are unknown. This study provides the first assessment of their biodiversity value by examining aspects of epigeal spider diversity supported by these non-target habitat patches in comparison to calcareous grassland. Spiders were sampled in each habitat from April to August 2014 using pitfall traps across three upland regions in Great Britain. Spider species assemblages were distinct between limestone heath and both grassland types. Distinction in species assemblages are likely due to differences in vegetation structure and microclimate, e.g., humidity, degree of shade. Each habitat type supported several rare species (e.g., Jacksonella falconeri, Agyneta subtilis) revealing the contribution to spider fauna. The distinct spider species assemblage and presence of rare species in limestone heath patches demonstrate their importance in the upland calcareous grassland matrix. This study highlights the value of monitoring biodiversity in non-target habitats within a habitat matrix alongside those that are actively targeted by management.     

Can long-distance dispersal shape the local and regional species pool?

Long-distance dispersal events are irregular and their role in shaping plant diversity is often discussed and modeled but rarely studied experimentally. We mimicked long-distance dispersal experimentally by sowing eleven exotic and fourteen native species into a calcareous grassland community in Estonia. Exotic species were randomly chosen from the collection of 500 herbaceous species in the Botanical Garden of the Tartu University. All exotic species were able to complete their life-cycles under the climatic and edaphic conditions in the garden. Native species originated from open dry calcareous habitats in the surroundings of the study site, but did not occur in the experimental grassland. Seven exotic species and seven native species established during the first year. In the third year, there were still three exotic species with five premature individuals, and three sown native species with sixteen individuals in the plots. These results show that long-distance dispersal both within and between regions may have an impact on species composition in target plant communities. If relatively the best established exotic speciesPhyteuma scheuchzeri would be classified as casual, one may conclude that transition among introduction and casual stages corresponds to ten’ rule. The species richness of seedlings, taking both local and sown species into account, was higher in plots with higher native established plant species richness.     

Process‐based long‐term evaluation of an ecological network of calcareous grasslands connected by sheep herding

This paper evaluates the long‐term effect of an ecological network of calcareous grasslands, a habitat type that experienced dramatic habitat loss and fragmentation during the 20th century, on species richness of habitat specialist plants. Calcareous grasslands are of special conservation concern as the habitat type with the highest diversity in plant and invertebrate species in central Europe. A baseline survey in 1989 established complete vascular plant species lists for all 62 previously abandoned calcareous grassland patches in the study area and assessed the presence of 48 habitat specialist plant species. An ecological network was initiated in 1989 to reconnect these patches with existing grazed pastures (core areas) through large flock sheep herding where feasible, as sheep are thought to be the primary dispersal vectors for calcareous grassland plants. An evaluation survey in 2009 showed significant increase in species richness of habitat specialist plants in patches reconnected by sheep herding, indicating successful colonizations by habitat specialist plants, while ungrazed patches showed no significant change. Observed increase in species richness between 1989 and 2009 was related to connectivity by sheep herding and the presence of a diversity of structural elements providing microsites for establishment. Baseline species richness of the patches, which had been abandoned since at least 1960, was associated with patch area, supporting the effect of ecological drift, and with vegetation type, which suggests that delays in extinction may be related to site factors governing the strength of competition with later seral species. The implementation of this ecological network represents a long‐term ‘natural experiment’ with baseline data, manipulation, and evaluation of hypothesized effects on a clearly defined target variable. It thus provides much needed empirical evidence that species loss in fragmented calcareous grassland communities can be counteracted by restoring functional connectivity among remnant patches.     

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.     

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.     

The influence of wild boar <Emphasis Type="Italic">(Sus scrofa)</Emphasis> on microhabitat quality for the endangered butterfly <Emphasis Type="Italic">Pyrgus malvae</Emphasis> in the Netherlands

The decline of open habitats in Europe, such as semi-natural grasslands and heathlands, has caused a general decline in biodiversity, which has been well documented for butterflies. Current conservation practices often involve grazing by domestic livestock to maintain suitable butterfly habitats. The extent to which wild ungulates may play a similar role remains largely unknown. Through their rooting activity, wild boar could be effective to reduce grass encroachment and restore pioneer microhabitats that are vital to many grassland insects in temperate climates. Here, we assessed the microhabitat requirements of Pyrgus malvae, an endangered butterfly of heathland and grassland habitats in the Netherlands, with special attention for the influence of wild boar rooting. To date, oviposition site selection of this species has concentrated on calcareous grasslands, whereas we also include heathlands. Overall, larval occupancy was higher in warm, open and sparsely vegetated microhabitats, which supports earlier findings. In heathland, microhabitat occupancy was positively affected by bryophyte and litter cover. In heath-grassland mosaic, microhabitat occupancy was also influenced by bryophyte and litter cover, but in addition low grass cover increased occupancy by favouring host plants. In grassland, only low grass cover and host plant cover determined microhabitat quality. Across all habitats, occupied microhabitats were characterized by lower vegetation as well as higher average daytime temperatures than unoccupied microhabitats. We discovered that wild boar play an important role in reducing grass cover by shallow rooting in grass patches, thereby increasing host plant availability. Hence, wild boar may have an added value in maintaining and restoring P. malvae microhabitats in grassland habitats in addition to grazing by domestic livestock.     

Effects of Habitat Quality and Isolation on the Colonization of Restored Heathlands by Butterflies

In areas with intensive land use, such as the Netherlands, habitat fragmentation and loss of habitat quality due to eutrophication and drainage are major threats to the preservation of species‐rich communities of heathland and acid grassland. Restoration of such nutrient‐poor habitats may be carried out by removing the topsoil from ex‐arable land, in order to lower the nutrient levels. However, the establishment of target plant communities is known to be fragmentary. The current study shows that this also applies to butterflies. Ten years after topsoil removal in eight study areas, on average, only 3.5 of 10 characteristic heathland species were recorded on the sites. Species that did colonize had a significantly lower density than in the source populations. Our study indicates that although isolation effects were limiting colonization, poor habitat quality was the main limiting factor, mainly due to lack of host plants, hydrological conditions, and, to a lesser extent, lack of nectar plants and excessive residual nutrient levels. An experiment with the introduction of cut heather in one study area showed a significantly higher abundance of both target and nontarget butterflies in manipulated sites than in control sites. It can be concluded that habitat restoration by topsoil removal can be successful for butterflies of especially wet heathland habitats, provided that source populations are at close range and care is taken that complete plant communities are restored.     

The potential bias of nitrogen deposition effects on primary productivity and biodiversity

Atmospheric nitrogen (N) deposition is composed of both inorganic nitrogen (IN) and organic nitrogen (ON), and these sources of N may exhibit different impacts on ecosystems. However, our understanding of the impacts of N deposition is largely based on experimental gradients of INs or more rarely ONs. Thus, the effects of N deposition on ecosystem productivity and biodiversity may be biased. We explored the differential impacts of N addition with different IN:ON ratios (0:10, 3:7, 5:5, 7:3, and 10:0) on aboveground net primary productivity (ANPP) of plant community and plant diversity in a typical temperate grassland with a long-term N addition experiment. Soil pH, litter biomass, soil IN concentration, and light penetration were measured to examine the potential mechanisms underlying species loss with N addition. Our results showed that N addition significantly increased plant community ANPP by 68.33%–105.50% and reduced species richness by 16.20%–37.99%. The IN:ON ratios showed no significant effects on plant community ANPP. However, IN-induced species richness loss was about 2.34 times of ON-induced richness loss. Soil pH was positively related to species richness, and they exhibited very similar response patterns to IN:ON ratios. It implies that soil acidification accounts for the different magnitudes of species loss with IN and ON additions. Overall, our study suggests that it might be reasonable to evaluate the effects of N deposition on plant community ANPP with either IN or ON addition. However, the evaluation of N deposition on biodiversity might be overestimated if only IN is added or underestimated if only ON is added.     

Atmospheric nitrogen deposition explains patterns of plant species loss

Atmospheric nitrogen (N) deposition across Europe increased substantially from the 1950s to the 1990s. Targeted surveys suggest a negative correlation between N deposition and species richness within quadrats in sensitive habitats. However, it remains unclear whether plant species losses at national recording scales are correlated with nitrogen deposition. We relate plant species losses before 1987 in Great Britain to reduced and oxidized N deposition, land use change and climate change. The mean Ellenberg fertility (N) indices of plant species lost in each 100 km² cell before 1987 was compared with those of species that were recorded between 1987 and 1999. In 45% of squares, indices of species lost were significantly lower than those for species present after 1986. For 17%, primarily upland, squares, the opposite effect was found. A generalized least squares regression model, with difference in the mean Ellenberg N index between samples as the dependent variable, showed that higher deposition of reduced N was significantly associated with selective loss of species with a lower index. Arable land use and change in arable land use also demonstrated this positive relationship. Rough grazing, change in rough grazing, change in pasture and change in annual precipitation showed negative effects. Difference in Ellenberg R index was highly correlated with difference in Ellenberg N and was negatively correlated with oxidized N deposition, suggesting that the lack of a significant effect of oxidized N deposition on Ellenberg N was because it had effects through both acidification and eutrophication, while the effect of reduced N deposition was primarily through eutrophication. Our results suggest that N deposition, along with land use and precipitation changes, has been a significant driver of local plant extinctions. With N deposition increasing in many parts of the world, local extinctions of plant species may be experienced in other regions.     

Impacts of atmospheric nitrogen deposition: responses of multiple plant and soil parameters across contrasting ecosystems in long‐term field experiments

Atmospheric nitrogen (N) deposition is a global and increasing threat to biodiversity and ecosystem function. Much of our current understanding of N deposition impacts comes from field manipulation studies, although interpretation may need caution where simulations of N deposition (in terms of dose, application rate and N form) have limited realism. Here, we review responses to simulated N deposition from the UKREATE network, a group of nine experimental sites across the UK in a diversity of heathland, grassland, bog and dune ecosystems which include studies with a high level of realism and where many are also the longest running globally on their ecosystem type. Clear responses were seen across the sites with the greatest sensitivity shown in cover and species richness of bryophytes and lichens. Productivity was also increased at sites where N was the limiting nutrient, while flowering also showed high sensitivity, with increases and declines seen in dominant shrub and forb species, respectively. Critically, these parameters were responsive to some of the lowest additional loadings of N (7.7–10 kg ha^?1 yr^?1) showing potential for impacts by deposition rates seen in even remote and ‘unpolluted’ regions of Europe. Other parameters were less sensitive, but nevertheless showed response to higher doses. These included increases in soil %N and ‘plant available’ KCl extractable N, N cycling rates and acid–base status. Furthermore, an analysis of accumulated dose that quantified response against the total N input over time suggested that N impacts can ‘build up’ within an ecosystem such that even relatively low N deposition rates can result in ecological responses if continued for long enough. Given the responses have important implications for ecosystem structure, function, and recovery from N loading, the clear evidence for impacts at relatively low N deposition rates across a wide range of habitats is of considerable concern.     

Effects of water and nitrogen addition on species turnover in temperate grasslands in northern China

Global nitrogen (N) deposition and climate change have been identified as two of the most important causes of current plant diversity loss. However, temporal patterns of species turnover underlying diversity changes in response to changing precipitation regimes and atmospheric N deposition have received inadequate attention. We carried out a manipulation experiment in a steppe and an old-field in North China from 2005 to 2009, to test the hypothesis that water addition enhances plant species richness through increase in the rate of species gain and decrease in the rate of species loss, while N addition has opposite effects on species changes. Our results showed that water addition increased the rate of species gain in both the steppe and the old field but decreased the rates of species loss and turnover in the old field. In contrast, N addition increased the rates of species loss and turnover in the steppe but decreased the rate of species gain in the old field. The rate of species change was greater in the old field than in the steppe. Water interacted with N to affect species richness and species turnover, indicating that the impacts of N on semi-arid grasslands were largely mediated by water availability. The temporal stability of communities was negatively correlated with rates of species loss and turnover, suggesting that water addition might enhance, but N addition would reduce the compositional stability of grasslands. Experimental results support our initial hypothesis and demonstrate that water and N availabilities differed in the effects on rate of species change in the temperate grasslands, and these effects also depend on grassland types and/or land-use history. Species gain and loss together contribute to the dynamic change of species richness in semi-arid grasslands under future climate change.     

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.     

Loss of forb diversity in relation to nitrogen deposition in the UK: regional trends and potential controls

In this study we investigate the impact of nitrogen (N) deposition on the diversity of three different vegetation functional groups – forbs, grasses and mosses – using a field survey of acid grasslands across Great Britain. Our aim is to identify the vegetation types that are most vulnerable to enhanced N deposition, and to shed light on the mechanisms that may be driving N‐initiated species changes in the UK. Sixty‐eight randomly selected grasslands belonging to the UK National Vegetation Classification group U4 (Festuca ovina–Agrostis capillaris–Galium saxatile grassland) were studied along a gradient of atmospheric N deposition ranging from 6 to 36 kg N ha^?1 yr^?1. At each site, vegetation was surveyed and samples were taken from the topsoil and subsoil. Aboveground plant material was collected from three species: a forb, grass and moss. Both the species richness and cover of forbs declined strongly with increasing N deposition, from greater than eight species/20% cover per m² quadrat at low levels of N to fewer than two species/5% cover at the highest N deposition levels. Grasses showed a weak but significant decline in species richness, and a trend toward increasing cover with increasing N input. Mosses showed no trends in either species richness or cover. Most of the decline in plant species richness could be accounted for by the level of ammonium deposition. Soil KCl‐extractable ammonium concentration showed a significant positive correlation with N input, but there was no relationship between N deposition and extractable nitrate. In the soil O/A horizon, there was no relationship between N deposition and %N, and only a very weak positive relationship between the level of N deposition and the C : N ratio. Finally, in the vegetation, there was no relationship between N deposition and either shoot tissue N concentration or N : P ratio for any of the three reference species. Combining our regional survey with the results of published N‐addition experiments provides compelling evidence that there has been a significant decline in the species richness and cover of forbs across Great Britain, and that the primary cause is competition due to an increase in the cover of grasses in response to enhanced deposition of reactive N, primarily NH₄⁺.     

大气氮沉降影响草地植物物种多样性机制研究综述

大气氮沉降对草地生态系统结构和功能的影响已成为全球变化生物学研究重点。大气氮沉降导致草地群落物种多样性降低已成为全球普遍现象,但其生物学机制还不清楚,因此有必要系统梳理大气氮沉降对全球不同草地生态系统的研究结果,以便在氮沉降背景下为我国草地生态系统的研究和管理制定科学决策。系统综述了氮沉降降低草地群落物种多样性的可能机制,主要包括资源竞争排斥、群落更新限制、土壤酸化及其离子毒害、养分失衡、氮素本身的毒害、次生胁迫。氮沉降导致草地物种多样性降低是多种机制综合作用的结果,每种机制在不同时空具有不同的相对贡献。同时,与欧洲酸性土壤草地和美国高草草原相比,我国草地土壤类型和植被属性具有明显差异。因此,应根据我国草地生态系统的特征、不同植物功能利用养分策略,从土壤养分变化、根系养分吸收转运、叶片生理过程等方面的整合研究思路,探讨氮沉降影响我国草地群落物种多样性的生物学机制,为我国草地生态系统的科学管理提供理论依据。     

Nitrogen addition decreases seed germination in a temperate steppe

Seed germination and seedling establishment play an important role in driving the responses of plant community structure and function to global change. Nitrogen (N) deposition is one of the driving factors of global change, which often leads to a loss in species richness in grassland ecosystems. However, how seed germination responds to N addition remains unclear. A pot incubation test was conducted in a semi‐arid grassland in the Mongolian Plateau, Northern China, to investigate the effect of N addition (0, 5, 10, 20, 40, and 80 g N/m²) on seed germination from May to October 2016. Twenty species germinated under all treatments; however, the responses of the 20 species to N addition were different. The densities of Stipa krylovii, Leymus chinensis, and Artemisia frigida, which are the dominant species in this temperate steppe, decreased significantly as the amount of N addition. Moreover, N addition significantly suppressed seedling densities of the community, perennial forbs, perennial grasses, and annuals and biennials. Furthermore, species richness of the community, perennial forbs, and annuals and biennials decreased sharply with increasing N addition level, but perennial grass species richness did not change. The Shannon–Wiener diversity index also decreased as the amount of N addition increased. Our results suggest that N enrichment plays an important role in the seed germination stage and decreases supplements of seedlings to adult plants. These findings may help explain the causes of species loss by atmospheric N deposition in grassland ecosystems.