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。     

Nutrient availability controls the impact of mammalian herbivores on soil carbon and nitrogen pools in grasslands

Grasslands are subject to considerable alteration due to human activities globally, including widespread changes in populations and composition of large mammalian herbivores and elevated supply of nutrients. Grassland soils remain important reservoirs of carbon (C) and nitrogen (N). Herbivores may affect both C and N pools and these changes likely interact with increases in soil nutrient availability. Given the scale of grassland soil fluxes, such changes can have striking consequences for atmospheric C concentrations and the climate. Here, we use the Nutrient Network experiment to examine the responses of soil C and N pools to mammalian herbivore exclusion across 22 grasslands, under ambient and elevated nutrient availabilities (fertilized with NPK + micronutrients). We show that the impact of herbivore exclusion on soil C and N pools depends on fertilization. Under ambient nutrient conditions, we observed no effect of herbivore exclusion, but under elevated nutrient supply, pools are smaller upon herbivore exclusion. The highest mean soil C and N pools were found in grazed and fertilized plots. The decrease in soil C and N upon herbivore exclusion in combination with fertilization correlated with a decrease in aboveground plant biomass and microbial activity, indicating a reduced storage of organic matter and microbial residues as soil C and N. The response of soil C and N pools to herbivore exclusion was contingent on temperature – herbivores likely cause losses of C and N in colder sites and increases in warmer sites. Additionally, grasslands that contain mammalian herbivores have the potential to sequester more N under increased temperature variability and nutrient enrichment than ungrazed grasslands. Our study highlights the importance of conserving mammalian herbivore populations in grasslands worldwide. We need to incorporate local‐scale herbivory, and its interaction with nutrient enrichment and climate, within global‐scale models to better predict land–atmosphere interactions under future climate change.     

Cumulative nitrogen input drives species loss in terrestrial ecosystems

Aim Elevated inputs of biologically reactive nitrogen (N) are considered to be one of the most substantial threats to biodiversity in terrestrial ecosystems. Several attempts have been made to scrutinize the factors driving species loss following excess N input, but generalizations across sites or vegetation types cannot yet be made. Here we focus on the relative importance of the vegetation type, the local environment (climate, soil pH, wet deposition load) and the experimentally applied (cumulative) N dose on the response of the vegetation to N addition. Location Mainly North America and Europe. Methods We conducted a large‐scale meta‐analysis of in situ N addition experiments in different vegetation types, focusing on the response of biomass and species richness. Results Whereas the biomass of grasslands and salt marshes significantly increased with N fertilization, forest understorey vegetation, heathlands, freshwater wetlands and bogs did not show any significant response. Graminoids significantly increased in biomass following N addition, whereas bryophytes significantly lost biomass; shrubs, forbs and lichens did not significantly respond. The yearly N fertilization dose significantly influenced the biomass response of grassland and salt marshes, while for the other vegetation types none of the collected predictor variables were of significant influence. Species richness significantly decreased with N addition in grasslands and heathlands [Correction added on 23 March 2011, after first online publication: ‘across all vegetation types’ changed to ‘in grasslands and heathlands’]. The relative change in species richness following N addition was significantly driven by the cumulative N dose. Main conclusions The decline in species richness with cumulative N input follows a negative exponential pathway. Species loss occurs faster at low levels of cumulative N input or at the beginning of the addition, followed by an increasingly slower species loss at higher cumulative N inputs. These findings lead us to stress the importance of including the cumulative effect of N additions in calculations of critical load values.     

Vegetation development after a large scale restoration of species-rich grasslands in a Central European floodplain

Large scale restoration using local high-diversity seed mixture combined with turf transfer was applied on ex-arable land in the Morava River floodplain in the western Slovakia in the years 1999–2012. The post-restoration vegetation development was recorded during 12 years after the restoration using floristic records per restored polygons with cover estimation in simple 3-degree scale. Temporal changes in species composition were evaluated by gradient analysis and number of characteristic grassland and ruderal species on restored sites was analysed by general linear models. Species composition changed gradually towards the species composition typical for species-rich floodplain grasslands, but the trajectory was not straightforward and several irregularities were observed. They were probably induced by extreme weather events (drought, floods). The decrease in ruderal species and increase in the number of typical floodplain grassland species were observed, when floodplain grassland species permanently outcompeted ruderal species since 8th year after the restoration. However the development in large scale was slower, than expected from previous small-scale experiments, it is evident, that combination of local seed mixture sowing with a turf transfer is a feasible method for the restoration of species-rich floodplain grasslands from arable land.     

Carbon fluxes and species diversity in grazed and fenced typical steppe grassland of Inner Mongolia,China

Aim Grasslands are dominant vegetation of China, support outstanding biodiversity and sequester bulk amount of atmospheric CO₂. These grasslands are highly degraded and fragmented due to remarkable anthropogenic and grazing loads. Chinese Government has made great attempt to restore by grazing exclusion. The relations of carbon fluxes with species composition and diversity in the communities sensitive to grazing by large herbivores are needed to be analysed under the global climate change scenario. The objective of present study was to comprehend the effects of grazing and fencing on the ecosystem structure and function of the typical steppe grassland.     

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.     

Changes in species composition of European acid grasslands observed along a gradient of nitrogen deposition

Question: Which environmental variables affect floristic species composition of acid grasslands in the Atlantic biogeographic region of Europe along a gradient of atmospheric N deposition? Location: Transect across the Atlantic biogeographic region of Europe including Ireland, Great Britain, Isle of Man, France, Belgium, The Netherlands, Germany, Norway, Denmark and Sweden. Materials and Methods: In 153 acid grasslands we assessed plant and bryophyte species composition, soil chemistry (pH, base cations, metals, nitrate and ammonium concentrations, total C and N, and Olsen plant available phosphorus), climatic variables, N deposition and S deposition. Ordination and variation partitioning were used to determine the relative importance of different drivers on the species composition of the studied grasslands. Results: Climate, soil and deposition variables explained 24% of the total variation in species composition. Variance partitioning showed that soil variables explained the most variation in the data set and that climate and geographic variables accounted for slightly less variation. Deposition variables (N and S deposition) explained 9.8% of the variation in the ordination. Species positively associated with N deposition included Holcus mollis and Leontodon hispidus. Species negatively associated with N deposition included Agrostis curtisii, Leontodon autumnalis, Campanula rotundifolia and Hylocomium splendens. Conclusion: Although secondary to climate gradients and soil biogeochemistry, and not as strong as for species richness, the impact of N and S deposition on species composition can be detected in acid grasslands, influencing community composition both directly and indirectly, presumably through soil‐mediated effects.     

Effects of enhanced nitrogen deposition and phosphorus limitation on nitrogen budgets of semi-natural grasslands

Increased reactive atmospheric N deposition has been implicated in floristic changes in species‐rich acidic and calcareous grasslands, but the fate of this pollutant N in these ecosystems is unknown. This paper reports the first analysis of N budgets and N fluxes for two grasslands in the White Peak area of Derbyshire, one of the most heavily N‐polluted locations in the UK. N fluxes were monitored in lysimeter cores (retaining the original turfs) taken from field plots of unimproved acidic and calcareous grasslands that had received (in addition to ambient N deposition) simulated enhanced N deposition treatments of 3.5 and 14 g N m^?2 yr^?1 for 6 years. The influence of reducing phosphorus limitation was assessed by factorial additions of P. Seasonal leached losses of nitrate, ammonia and organic N were monitored in detail along with estimates of N removal through simulated grazing and gaseous losses through denitrification and volatilization. The rates of N fluxes by these pathways were used to create N budgets for the grasslands. Both grasslands were found to be accumulating much of the simulated additional N deposition: up to 89% accumulated in the calcareous grassland and up to 38% accumulated in the acidic grassland. The major fluxes of N loss from these grasslands were by simulated grazing and leaching of soluble organic N (constituting 90% of leached N under ambient conditions). Leached inorganic N (mainly nitrate) contributed significantly to the output flux of N under the highest N treatment only. Loss of N through ammonia volatilization accounted for less than 6% of the N added as simulated deposition, while denitrification contributed significantly to output fluxes only in the acidic grassland during winter. The implications of the results for ecosystem N balances and the likely consequences of N accumulation on these grasslands are discussed.     

Future changes and driving factors of global peak vegetation growth based on CMIP6 simulations

Accurate detection and attribution of changes in global peak vegetation growth at the annual scale are prerequisites for characterising the productivity of terrestrial ecosystems and developing strategies for the sustainable management of ecosystems. This study examined the long-term global normalised difference vegetation index during the baseline period (1982–2015) and found widespread greening in 70% of global vegetated areas in response to climate warming. However, climate change is not the only cause of global greening. The spatial variability in the response of global vegetation to environmental factors has not been well established. The Cubist model was used to investigate the relationship between peak vegetation growth and environmental variables. The results showed that 64% of the spatial variation in greening/browning can be explained by climate (including precipitation and temperature), followed by atmospheric components of nitrogen deposition and carbon dioxide concentration (17%), terrain properties (12%), and soil properties (7%). By incorporating future climate and atmospheric component projections from the Coupled Model Intercomparison Project Phase 6 into the model, enhanced vegetation greening was predicted globally, particularly in evergreen needle-leaf forests and grasslands, from 2081 to 2100. Many browning changes were predicted in evergreen and deciduous broadleaf forests, mixed forests, and around areas influenced by human land use. Overall, these findings reveal that environmental factors have relevant integrated impacts on vegetation dynamics under climate change and should be considered during the design of local mitigation and adaptation management strategies.     

Recent changes in mountain grasslands: a vegetation resampling study

Understanding how land‐use changes affect different facets of plant biodiversity in seminatural European grasslands is of particular importance for biodiversity conservation. As conclusions of previous experimental or synchronic observational studies did not converge toward a general agreement, assessing the recent trends in vegetation change in various grassland systems using a diachronic approach is needed. In this resurvey study, we investigated the recent changes in grassland vegetation of the French Jura Mountains, a region with a long tradition of pastoralism. We compared the floristic composition of 150 grassland plots recorded between 1990 and 2000 with new relevés made in 2012 on the same plots. We considered taxonomic, phylogenetic and functional diversity as well as ecological characteristics of the plant communities derived from ecological indicator values and life strategies of the species. PCA of the floristic composition revealed a significant general trend linked to the sampling year. Wilcoxon paired tests showed that contemporary communities were generally more dominated by grass species and presented a higher tolerance to defoliation, a higher pastoral value, and a higher nutrient indicator value. Comparisons revealed a decrease in phylogenetic and functional diversity. By contrast, local species richness has slightly increased. The intensity of change in species composition, measured by Hellinger distance between pairs of relevés, was dependent on neither the time lag between the two surveys, the author of the first relevé nor its location or elevation. The most important changes were observed in grasslands that previously presented low pastoral value, low grass cover, low tolerance to defoliation, and high proportion of stress‐tolerant species. This trend was likely linked to the intensification of grassland management reported in the region, with a parallel increase in mowing frequency, grazing pressure, and fertilization level. More restrictive specifications should be applied to agricultural practices to avoid overexploitation of mountain species‐rich grasslands and its negative consequences on their biodiversity and resilience.     

Future climate change effects on US forest composition may offset benefits of reduced atmospheric deposition of N and S

Climate change and atmospheric deposition of nitrogen (N) and sulfur (S) are important drivers of forest demography. Here we apply previously derived growth and survival responses for 94 tree species, representing >90% of the contiguous US forest basal area, to project how changes in mean annual temperature, precipitation, and N and S deposition from 20 different future scenarios may affect forest composition to 2100. We find that under the low climate change scenario (RCP 4.5), reductions in aboveground tree biomass from higher temperatures are roughly offset by increases in aboveground tree biomass from reductions in N and S deposition. However, under the higher climate change scenario (RCP 8.5) the decreases from climate change overwhelm increases from reductions in N and S deposition. These broad trends underlie wide variation among species. We found averaged across temperature scenarios the relative abundance of 60 species were projected to decrease more than 5% and 20 species were projected to increase more than 5%; and reductions of N and S deposition led to a decrease for 13 species and an increase for 40 species. This suggests large shifts in the composition of US forests in the future. Negative climate effects were mostly from elevated temperature and were not offset by scenarios with wetter conditions. We found that by 2100 an estimated 1 billion trees under the RCP 4.5 scenario and 20 billion trees under the RCP 8.5 scenario may be pushed outside the temperature record upon which these relationships were derived. These results may not fully capture future changes in forest composition as several other factors were not included. Overall efforts to reduce atmospheric deposition of N and S will likely be insufficient to overcome climate change impacts on forest demography across much of the United States unless we adhere to the low climate change scenario.     

Livestock grazing affects microclimate conditions for decomposition process through changes in vegetation structure in mountain grasslands

It is often assumed that a change in litter quality is the main driver of alterations in the decomposition process when grazers modify vegetation structure. Soil microclimate is also modified, but this driver of decomposition has been far less studied than litter quality. We analyzed the relationships among vegetation structure, microclimate and decomposition in different mountain grassland types, across a fence-line separating paddocks with different grazing intensity. Along the fence, we selected nine pairs of contrasting grassland types including lawns and tall tussock grasslands, which are associated with high and low local grazing pressure, respectively. At each site (N = 18) we estimated growth form composition and vegetation height. During the growing season we recorded soil temperature, soil moisture and the photosynthetically active radiation. Within the same period, we measured the decomposition rate of two common litter substrates. We analyzed the relationships among those variables at the landscape and at the local scale. At the landscape scale we considered the variation across all sites (N = 18). At the local scale we considered each pair as a sample (N = 9) and the differences between both sides of the fence as the variables to correlate. Our results indicate that when short grasslands are released from grazing and tall grasslands became dominant, temperature and light at the soil level are reduced, while soil moisture tends to increase, enhancing decomposition. Furthermore, these results show that the microclimatic conditions effect can counteract the litter quality effect (reported in previous studies) on decomposition, resulting in increased decomposition rates when grazing is reduced.     

Near‐natural methods promote restoration of species‐rich grassland vegetation—revisiting a road verge trial after 9 years

The present loss of species‐rich grasslands makes it vital to restore these valuable habitat types, including novel habitat variants such as road verges. Due to the lack of knowledge on long‐term outcomes of restoration initiatives, well‐designed studies comparing different restoration methods are needed. In this study, we examined fine‐scale vegetation recovery patterns over 9 years in a field experiment with several near‐natural restoration methods (adding local seed mixtures, transferring hay from local grasslands using hard or light raking, and natural regeneration) in a road verge. We compared this to standard revegetation (hydroseeding species‐poor commercial seed mixtures). We found major temporal changes in vegetation restored by local seed or hay transfer, before it gradually became more similar to the donor grasslands and seed mixtures, which served as references for the experiment. Natural (spontaneous) regeneration with seed dispersal from surroundings gave similar results, whereas areas revegetated using standard methods became more dissimilar to the reference sites during the study period. The main variation in species composition reflected the contrast between local donor grasslands and seed mixtures and the species‐poor early successional grasslands. We conclude that near‐natural methods (hay transfer and seeding) successfully restored species‐rich grassland, including road verges. This study underlines the importance of comparing several treatments over a sufficiently long period to assess their success in restoring species‐rich grassland.     

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.     

Bird community composition in an actively managed savanna reserve, importance of vegetation structure and vegetation composition

The mosaic of trees, shrubs and open grassland in mesic African savannas is highly dynamic and strongly influenced by mammal herbivory and fire. We investigated the bird fauna in four different savanna habitats to help assess the impacts of vegetation change on this component of faunal diversity. Birds were censused, plant species were identified and vegetation structure was measured in four different vegetation types (Acacia nilotica woodland, Acacia nigrescens woodland, broadleaf thicket and open grassland) in the Hluhluwe-Umfolozi Park in northern KwaZulu Natal, South Africa. Multivariate ordination analyses were used to determine the relative importance of vegetation structure and floristic composition in defining bird assemblages. The bird communities of the grasslands, the acacia woodlands, and the broadleaf woodlands were clearly separated on the first axis of the detrended canonical correspondence analysis (DCCA). Canopy cover and foliage height diversity (FHD) were strongly correlated with the first axis of DCCA, possibly reflecting a secondary successional series from grassland to woodland, known as bush encroachment. Floristic composition (based on presence–absence data only) seemed to be less important for bird community composition than vegetation structure. The results indicate that changes in vegetation structure, caused by bush encroachment, could cause concomitant changes in bird community composition.     

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.     

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.     

A global comparison of grassland biomass responses to CO2 and nitrogen enrichment

Grassland ecosystems cover vast areas of the Earth''s surface and provide many ecosystem services including carbon (C) storage, biodiversity preservation and the production of livestock forage. Predicting the future delivery of these services is difficult, because widespread changes in atmospheric CO₂ concentration, climate and nitrogen (N) inputs are expected. We compiled published data from global change driver manipulation experiments and combined these with climate data to assess grassland biomass responses to CO₂ and N enrichment across a range of climates. CO₂ and N enrichment generally increased aboveground biomass (AGB) but effects of CO₂ enrichment were weaker than those of N. The response to N was also dependent on the amount of N added and rainfall, with a greater response in high precipitation regions. No relationship between response to CO₂ and climate was detected within our dataset, thus suggesting that other site characteristics, e.g. soils and plant community composition, are more important regulators of grassland responses to CO₂. A statistical model of AGB response to N was used in conjunction with projected N deposition data to estimate changes to future biomass stocks. This highlighted several potential hotspots (e.g. in some regions of China and India) of grassland AGB gain. Possible benefits for C sequestration and forage production in these regions may be offset by declines in plant biodiversity caused by these biomass gains, thus necessitating careful management if ecosystem service delivery is to be maximized. An approach such as ours, in which meta-analysis is combined with global scale model outputs to make large-scale predictions, may complement the results of dynamic global vegetation models, thus allowing us to form better predictions of biosphere responses to environmental change.     

Scale‐dependence of vegetation‐environment relationships in semi‐natural grasslands

Questions: Which environmental and management factors determine plant species composition in semi‐natural grasslands within a local study area? Are vegetation and explanatory factors scale‐dependent? Location: Semi‐natural grasslands in Lærdal, Sognog Fjordane County, western Norway. Methods: We recorded plant species composition and explanatory variables in six grassland sites using a hierarchically nested sampling design with three levels: plots randomly placed within blocks selected within sites. We evaluated vegetation‐environment relationships at all three levels by means of DCA ordination and split‐plot GLM analyses. Results: The most important complex gradient determining variation in grassland species composition showed a broad‐scale relationship with management. Soil moisture conditions were related to vegetation variation on block scale, whereas element concentrations in the soil were significantly related to variation in species composition on all spatial scales. Our results show that vegetation‐environment relationships are dependent on the scale of observation. We suggest that scale‐related (and therefore methodological) issues may explain the wide range of vegetation‐environment relationships reported in the literature, for semi‐natural grassland in particular but also for other ecosystems. Conclusions: Interpretation of the variation in species composition of semi‐natural grasslands requires consideration of the spatial scales on which important environmental variables vary.     

Plant responsiveness to variation in precipitation and nitrogen is consistent across the compositional diversity of a California annual grassland

Question: How does responsiveness to water and Nitrogen (N) availability vary across the compositional and functional diversity that exists in a mesic California annual grassland plant community? Location: Northern California annual grassland. Methods: A mesocosm system was used to simulate average annual precipitation totals and dry and wet year extremes observed in northern California mesic grasslands. The effects of precipitation and N availability on biomass and fecundity were measured on three different vegetation types, a mixed grass forb community, and a forb and a grass monoculture. The treatment effects on plant community composition were examined in the mixed species community. Results: While growth and seed production of the three vegetation types was inherently different, their responses to variation in precipitation and N were statistically similar. Plant density, shoot biomass, and seed production tended to increase with greater water availability in all vegetation types, with the exception of a consistent growth reduction in high precipitation (1245 mm) plots in the first year of the study. Shoot biomass responded positively to N addition, an effect that increased with greater water availability. Nitrogen addition had little effect on plant density or seed production. In the mixed grass‐forb community, biomass responsiveness to water and N treatments were consistently driven by the shoot growth of Avena barbata, the dominant grass species. Conclusions: Vegetation responses to changes in precipitation and N availability were consistent across a range of composition and structural diversity in this study. Plant growth and seed production were sensitive to both increased and decreased precipitation totals, and the magnitude of these responses to N availability varied depending on soil moisture conditions. Our results suggest the impacts of changing precipitation regimes and N deposition on annual productivity of California grasslands may be predictable under different climate scenarios across a range of plant communities.