Species richness loss after nutrient addition as affected by N:C ratios and phytohormone GA3 contents in an alpine meadow community

Aims From the light-competition hypothesis, competition for light is asymmetric and the observed increases in plant-size variability with increasingly denser canopies are primarily due to competition for light. Greater plant height provides pre-emptive access to light and produces increased height differences among species. The question is what produces these differences in plant height or height growth response among species in response to fertilization.Methods In 2009, a field experiment of N, P and N + P enrichments at three levels each was initiated in an alpine meadow on the northeast Qinghai-Tibet Plateau. Effects of fertilization on species richness, aboveground net primary production (ANPP), relative light intensity and plant height of different plant functional groups were determined. Festuca ovina (grass), Kobresia humilis (sedge), Oxytropis ochrocephala (legume), Taraxacum lugubre (rosette forb) and Geranium pylzowianum (upright forb) were selected as exemplars of each of the indicated functional groups. The N:C ratios in aboveground biomass, gibberellic acid (GA 3) concentrations in leaves, plant heights and height relative growth rate (RGR) of these exemplar species were analyzed in detail.Important findings Species richness of grasses significantly increased with increasing N + P levels. Species richness of legumes and upright forbs decreased after N and N + P additions. P addition had no significant effect on species richness. The effects of N + P addition on species richness and ANPP were consistently stronger than those of the single N or P fertilization. Reductions in species richness caused by nutrient addition paralleled the increases in ANPP and decreases in light intensity under the canopies, indicating indirect effect of nutrient addition on species richness via ANPP-induced light competition. The exemplar species that responded most positively to fertilization in height and RGR also displayed stronger increases in their GA 3 content and N:C ratios. GA 3 concentrations and N:C ratios were positively correlated with height RGR when the data were pooled for all species. The tallest and the fastest-growing grass, F. ovina, had the largest increase in N:C ratios and the highest leaf GA 3 concentrations after nutrient addition. These results indicated that differential responses of GA 3 concentrations and N:C ratios to fertilization were related to the inequality in plant heights among species.     

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.     

养分添加对天山高寒草地植物多样性和地上生物量的影响

为揭示高寒草地物种多样性和地上生物量以及二者之间关系对养分添加的响应模式, 该研究以天山高寒草地为对象, 通过两年的多重养分添加实验, 研究氮(N)、磷(P)、钾(K) 3种养分单独和组合添加对天山高寒草地群落物种多样性和地上生物量的影响。结果表明: (1)养分添加使当地植物物种多样性不同程度地减少, 其中以N + P、N + K、N + P + K添加的效应最为显著, 多重养分添加导致的土壤生态位维度降低是当地物种丧失的重要原因。(2)养分添加能显著提高群落地上生物量, 其中N为第一限制养分, 解除N限制后P和K成为限制养分, N + P + K复合添加对地上生物量的提高最为显著。(3)养分添加两年后, 地上生物量与物种丰富度之间无显著回归关系且地上生物量增加主要是由于禾草类生物量增加导致, 说明地上生物量主要由少数优势种决定而非群落物种数。     

Plant community and tissue chemistry responses to fertilizer and litter nutrient manipulations in a temperate grassland

Human-mediated nutrient amendments have widespread effects on plant communities. One of the major consequences has been the loss of species diversity under increased nutrient inputs. The loss of species can be functional group dependent with certain functional groups being more prone to decline than others. We present results from the sixth year of a long-term fertilization and litter manipulation study in an old-field grassland. We measured plant tissue chemistry (C:N ratio) to understand the role of plant physiological responses in the increase or decline of functional groups under nutrient manipulations. Fertilized plots had significantly more total aboveground biomass and live biomass than unfertilized plots, which was largely due to greater productivity by exotic C₃ grasses. We found that both fertilization and litter treatments affected plant species richness. Species richness was lower on plots that were fertilized or had litter intact; species losses were primarily from forbs and non-Poaceae graminoids. C₃ grasses and forbs had lower C:N ratios under fertilization with forbs having marginally greater %N responses to fertilization than grasses. Tissue chemistry in the C₃ grasses also varied depending on tissue type with reproductive tillers having higher C:N ratios than vegetative tillers. Although forbs had greater tissue chemistry responses to fertilization, they did not have a similar positive response in productivity and the number of forb species is decreasing on our experimental plots. Overall, differential nutrient uptake and use among functional groups influenced biomass production and species interactions, favoring exotic C₃ grasses and leading to their dominance. These data suggest functional groups may differ in their responses to anthropogenic nutrient amendments, ultimately influencing plant community composition.     

Vegetation dynamics following resource manipulations in herb-rich woodland

Changes in species richness, turnover, composition and above-ground biomass of herb-rich woodland were documented following fertilizer application and water addition over three growing seasons. Addition of fertilizer significantly reduced species richness relative to unmanipulated control and water addition plots after 3 years. This change coincided with significant increases in biomass, which were largely due to increased growth of exotic annual grasses. The reductions in richness observed in the fertilized plots were a consequence of both lower rates of local colonization and enhanced rates of local extinction of the resident species. Species loss was not random; native species were lost after nutrient addition whereas exotic species were not. Nutrient limitation was more important for species coexistence in these communities than was water availability.     

Short-term responses of an alpine meadow community to removal of a dominant species along a fertilization gradient

Aims The relationship between biodiversity and ecosystem functioning has intrigued ecologists for several decades, but the effect of loss of a dominant species on community structure and functioning along a nutrient gradient remains poorly understood. The aim of this paper was to test the effect of a dominant species on community structure and function by conducting a species removal experiment along a fertilization gradient.Methods We removed the population of a dominant species (Elymus nutans) in a long-term fertilization field in an alpine meadow on the Tibetan Plateau, China. Univariate general linear models were used to evaluate the effects of fertilization and removal on above-ground vegetation characteristics, including photosynthetically active radiation in the understory, species richness, Shannon–Weiner diversity index, Simpson's dominance index, above-ground biomass (including different functional groups) and seedling richness and density.Important findings Results revealed that after two plant growing seasons, there was no significant effect of the removal of a dominant species on species richness and diversity of the remaining vegetation, but the biomass of forbs and seedling recruitment were significantly increased indicative of the potential for long-term effects. Moreover, removal had a large effect at high fertilization levels, but little effect when fertilization levels were low. Our studies indicated that community response to loss of a dominant species was mainly dependent on resource availability and the remaining functional group identities. We also found seedling recruitment was usually more sensitive to the influence of competition of dominant species than the established vegetation in the short term.     

The effects of fertilization on the trait-abundance relationships in a Tibetan alpine meadow community

Aims Comparisons of the trait–abundance relationships from various habitat types are critical for community ecology, which can offer us insights about the mechanisms underlying the local community assembly, such as the relative role of neutral vs. niche processes in shaping community structure. Here, we explored the responses of trait–abundance relationships to nitrogen (N), phosphorus (P) and potassium (K) fertilization in an alpine meadow.Methods Five fertilization treatments (an unfertilized control and additions of N, P, K and NPK respectively) were implemented using randomized block design in an alpine Tibetan meadow. Species relative abundance (SRA), plant above-ground biomass and species richness were measured in each plot. For 24 common species, we measured species functional traits: saturated height, specific leaf area (SLA) and leaf dry matter content (LDMC) in each treatment but seed size only in the unfertilized control. Standard major axis (SMA) regression and phylogenetically independent contrasts (PICs) analysis were used to analyse species trait–abundance relationships in response to different fertilization treatments.Important findings Positive correlations between SRA and saturated height were raised following N, P and NPK fertilizations, which indicated an increase in light competition in these plots. In P fertilized plots, SRA was also positively correlated with LDMC because tall grasses with a nutrients conservation strategy often have a relative competitive advantage in capturing limited light and soil nutrients. In K fertilized plots, neither the trait–abundance relationships nor above-ground biomass or species richness significantly differed from that in the control, which suggests that K was not a limiting resource in our study site. These significant correlations between species traits and relative abundance in fertilized treatment suggest that trait-based selection plays an important role in determining species abundance within local communities in alpine meadows.     

氮素添加对内蒙古退化草原生产力的短期影响

不合理的土地利用方式以及气候变化导致我国草原生态系统普遍退化, 主要表现在土壤养分降低、植被覆盖度减少、生产力下降。外源氮素添加是促进退化草原尽快恢复的一项重要措施, 尤其是对那些退化较为严重的草原。该研究选取内蒙古东乌珠穆沁旗不同退化程度(轻度、中度和重度)的草原群落, 于2014-2015年开展连续两年的氮素添加实验, 设置对照(不添加)、低水平(5.0 g N·m^-2·a^-1)、中水平(10.0 g N·m^-2·a^-1)和高水平(20.0 g N·m^-2·a^-1) 4种氮素添加处理, 探讨退化草原群落生产力在恢复过程中对不同水平氮素添加的响应。结果显示: (1)高、中水平氮素添加显著提高了轻度退化群落的地上生物量, 分别比对照增加了53.1%、51.6%, 氮素各水平添加对中度、重度群落地上生物量无显著影响; (2)高、中水平氮素添加显著提高了轻度退化群落中多年生根茎型禾草地上生物量, 分别比对照增加了45.1%、47.7%, 而多年生杂类草地上生物量分别比对照减少了37.4%、42.1%, 但中度和重度退化群落各功能群生物量的响应不显著; (3)三种水平氮素添加对轻、中、重度退化群落物种丰富度在试验期间均没有显著影响。研究结果表明氮素添加有助于提高轻度退化草原中多年生根茎型禾草的生物量, 进而提高群落的生物量, 但多年生杂类草会被逐渐替代, 导致生物量降低, 可见施氮对草原恢复的影响取决于草原退化 程度。     

Environmental and plant community determinants of species loss following nitrogen enrichment

Global energy use and food production have increased nitrogen inputs to ecosystems worldwide, impacting plant community diversity, composition, and function. Previous studies show considerable variation across terrestrial herbaceous ecosystems in the magnitude of species loss following nitrogen (N) enrichment. What controls this variation remains unknown. We present results from 23 N-addition experiments across North America, representing a range of climatic, soil and plant community properties, to determine conditions that lead to greater diversity decline. Species loss in these communities ranged from 0 to 65% of control richness. Using hierarchical structural equation modelling, we found greater species loss in communities with a lower soil cation exchange capacity, colder regional temperature, and larger production increase following N addition, independent of initial species richness, plant productivity, and the relative abundance of most plant functional groups. Our results indicate sensitivity to N addition is co-determined by environmental conditions and production responsiveness, which overwhelm the effects of initial community structure and composition.     

Influence of nitrogen addition on the primary production in Nei Mongol degraded grassland

Aims Irrational utilization and global climate change have caused degradation of grassland ecosystems in northern China with low soil fertility, decreased vegetation coverage and productivity. Nitrogen addition has been suggested an effective way to enhance restoration of those degraded grasslands. In this study, we selected a typical steppe with three different degrading levels, including lightly, moderately and heavily degraded communities, in East Ujimqin, Nei Mongol. Our objectives of this study are to examine if and how nitrogen (N) addition can enhance restoration of those degraded grasslands Methods Treatments with four levels of N addition (0, 5.0, 10.0 and 20.0 g N·m^-2·a^-1) were conducted to each of the three degraded communities from 2014 to 2015. Nitrogen was applied as urea in June of both years. Aboveground biomass was collected at the species level in 1 m × 1 m plot in August each year, all species biomass was summed as net primary production, and biomass of plant functional groups was calculated by perennial rhizome grasses, perennial bunchgrasses, perennial forbs, shrubs and semi-shrubs, annuals and biennials.Important findings Our results showed that the high (20.0 g N·m^-2·a^-1) and medium level N addition (10.0 g N·m^-2·a^-1) significantly increased the aboveground biomass of the slightly degraded community by 53.1% and 51.6% compared with no N addition. N addition had no significant effects on the moderately and heavily degraded communities. N addition with high and medium levels increased aboveground biomass of perennial rhizome grasses by 45.1% and 47.7%, but decreased that of perennial forbs by 37.4% and 42.1% at the slightly degraded community. Our results indicated that N addition could increase the growth of perennial rhizome grasses, and the growth of perennial forbs was suppressed consequently. Our results suggest that even the application of N fertilizers can only be helpful to restoration of those slightly degraded grasslands. Besides, N addition had no significant effects on species richness in different degraded communities indicating the fact that the study may not last long enough. For the purpose of increasing aboveground biomass of degraded grassland, we should not only consider the type and quantity of fertilization, but also the attribute of the degraded communities. In addition, the response of degraded community in biomass may strongly be impacted by degrading level of studied grassland.     

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⁺.     

氮素添加对科尔沁沙质草地物种多样性和生产力的影响

物种多样性和生产力是生态系统结构和功能的重要指标.以科尔沁沙质草地为对象,通过对其进行不同梯度的氮素添加处理,研究氮素添加对沙质草地生态系统物种组成、物种多样性和生产力的影响.结果表明：氮素添加改变了群落物种组成和群落中的优势种,使植被的高度和盖度增加,植被的透光率减小;随着氮素水平的增加,群落中物种丰富度减小,物种多样性降低;不同水平的氮素添加均显著增加了(P<0.01)群落地上生物量;物种丰富度与植被透光率呈线性正相关(P<0.01),与植被盖度呈线性负相关关系(P<0.01),说明长期的氮沉降与人为氮素输入将影响沙地生态系统的物种组成、物种多样性以及生产力.     

Plant species loss from European semi‐natural grasslands following nutrient enrichment – is it nitrogen or is it phosphorus?

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

草原化荒漠草本植物对人工施加磷素的响应

荒漠地区因土壤水分和养分含量较低而限制了植被的生长,且磷素对于植物的生长具有重要的作用。利用人工施加磷素的控制实验研究了不同磷肥处理对荒漠区草本植物物种丰富度、多度、盖度、生物量、植物株高等群落学特征的影响。实验表明：物种丰富度和多度在施肥量分别为12.5,25和50 g?m-2的梯度下,施肥当年和第二年相较于对照均有所降低,且施肥梯度越高,降低越明显;植被盖度和地上部生物量则在两年的实验中表现出相似的规律,在不同的施肥梯度下均有所提高,高肥处理对其促进作用更大,且在降水充足的07年高于降水较少的08年,说明水肥耦合更有助于群落生产力的提高;优势种株高则对磷素的响应存在种间差异,年际间的差异也较大,这或许与荒漠植物本身特有的生物学特性有关。     

Landscape-scale variation in the seed banks of floodplain wetlands with contrasting hydrology in China

1. At a local scale, the species composition, diversity and spatial variation of wetland plant communities are determined primarily by spatial and temporal heterogeneity in their environments. Less is known about variation at a landscape‐level. The floodplain of the Changjiang (Yangtze) River in China includes hydrologically connected, subtropical wetlands with different hydrological characteristics. 2. We examined seed‐bank species composition and richness in marshes of two contrasting hydrological types: permanent marshes, fed by local runoff, and lakeshore marshes more closely connected to the regulated river. Lakeshore marshes are flooded annually to depth of approximately 1 m and during flooding they support an alternate, aquatic vegetation type. The soil seed bank in March was a comparative estimator of species diversity. At the beginning of the growing season it included seeds from both phases of alternating vegetation types associated with the annual hydrological cycle. 3. A regional pool of 101 species was detected in the seed banks of six wetlands associated with the river and its tributaries: 56 occurred in permanent marshes and 59 in lakeshore marshes, with only 15 common to both. Species rarefaction curves indicated that more species occurred in permanent than lakeshore marshes at equal numbers of individuals sampled. However, the more heterogeneous lakeshore seed banks were estimated (Chao 2) to have greater total species richness (81) than permanent marsh (60). 4. Analysis using Sørensen's coefficient of similarity and DCA ordination revealed complex variation, with much greater differences between hydrological types than within them, irrespective of geographical distance. The types also differed significantly in the composition of four functional groups of species. 5. Despite the potential for dispersal of propagules via the annually pulsing river system (hydrochory), at a regional and landscape scale, diversity is maintained largely by large‐scale temporal hydrological heterogeneity and smaller scale spatial and topographic heterogeneity.     

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。     

不同类型高寒草地群落结构与生产对施氮的响应及其敏感性

大气氮沉降增加被认为是目前重要的环境问题,会引起生物多样性的丧失和生态系统稳定性的降低。但作为草地改良的管理措施,养分添加被广泛应用于退化草地的恢复。但由于不同类型草地所处气候与群落组成的差异,对氮输入的响应可能不同。通过在藏北高原高寒草甸与高寒草甸草原设定长期氮添加梯度试验(对照, 25, 50, 100, 200 kg N hm~(-2) a~(-1)),来探讨氮输入对生物多样性与生产的影响,并估算不同类型高寒草地的氮饱和阈值。施氮对高寒草甸物种多样性指数无影响,而随着施氮量的提高高寒草甸草原植物物种数和多样性指数均逐渐降低。开始施肥前两年,随着施氮量提高高寒草甸地上生物量呈现逐渐增加趋势,随着施肥时间的延长地上生物量呈现先增加后降低的趋势。在高寒草甸草原随着施氮量提高地上生物量均呈现先增加后降低的趋势。随着施氮量提高,开始施氮前三年高寒草甸禾草植物地上生物量逐渐提高;随着施氮时间的延长,禾草和豆科植物地上生物量呈现先增加后降低的趋势。高寒草甸莎草植物地上生物量由施氮开始时的逐渐增加转变为先增加后降低趋势,最后变为逐渐降低的趋势,这说明施氮不利于莎草植物的生长。施氮只在施肥第四年显著提高杂草植物地上生物量。高寒草甸草原呈现不同的规律,开始施氮前三年随着施氮量提高,禾草植物地上生物量呈现先增加后降低的趋势;随着施氮时间的延长,禾草地上生物量逐渐提高。莎草和杂草植物地上生物量呈现先增加后降低趋势。利用对氮输入响应最敏感的植物功能群禾草生物量估算的高寒草甸和高寒草甸草原的氮饱和阈值分别是109.5、125.8 kg N hm~(-2) a~(-1),这说明高寒草甸氮敏感性显著高于高寒草甸草原。由此可见,未来氮沉降增加会对不同类型高寒草地产生不同的影响,在不同类型高寒草地进行施肥恢复时也应将氮饱和阈值的差异考虑在内。     

Tradeoffs and thresholds in the effects of nitrogen addition on biodiversity and ecosystem functioning: evidence from inner Mongolia Grasslands

Nitrogen (N) deposition is widely considered an environmental problem that leads to biodiversity loss and reduced ecosystem resilience; but, N fertilization has also been used as a management tool for enhancing primary production and ground cover, thereby promoting the restoration of degraded lands. However, empirical evaluation of these contrasting impacts is lacking. We tested the dual effects of N enrichment on biodiversity and ecosystem functioning at different organizational levels (i.e., plant species, functional groups, and community) by adding N at 0, 1.75, 5.25, 10.5, 17.5, and 28.0 g N m^?2 yr^?1 for four years in two contrasting field sites in Inner Mongolia: an undisturbed mature grassland and a nearby degraded grassland of the same type. N addition had both quantitatively and qualitatively different effects on the two communities. In the mature community, N addition led to a large reduction in species richness, accompanied by increased dominance of early successional annuals and loss of perennial grasses and forbs at all N input rates. In the degraded community, however, N addition increased the productivity and dominance of perennial rhizomatous grasses, with only a slight reduction in species richness and no significant change in annual abundance. The mature grassland was much more sensitive to N‐induced changes in community structure, likely as a result of higher soil moisture accentuating limitation by N alone. Our findings suggest that the critical threshold for N‐induced species loss to mature Eurasian grasslands is below 1.75 g N m^?2 yr^?1, and that changes in aboveground biomass, species richness, and plant functional group composition to both mature and degraded ecosystems saturate at N addition rates of approximately 10.5 g N m^?2 yr^?1. This work highlights the tradeoffs that exist in assessing the total impact of N deposition on ecosystem function.     

Disentangling the role of management,vegetation structure,and plant quality for Orthoptera in lowland meadows

Seminatural grasslands provide habitats for various species and are important for biodiversity conservation. The understanding of the diverse responses of species and traits to different grassland managenient methods is therefore urgently needed. We disentangled the role of grassland management (fertilization and irrigation), vegetation structure (biomass, sward height) and plant quality (protein and fiber content) for Orthoptera communities in lowland hay meadows in Germany. We found vegetation structure to be the most important environmental category in explaining community structure of Orthoptera (species richness, total individuals, fiinctional diversity and species composition). Intensively used meadows (fertilized, irrigated, high plant biomass) were characterized by assemblages with few species, low functional diversity, and low conservation value. Thereby, the relatively moderate fertilizer inputs in our study system of up to -75 kg N/ha/year reduced functional diversity of Orthoptera, while this negative effect of fertilization was not detectable when solely considering taxonomic aspects. We found strong support for a prominent role of plant quality in shaping Orthoptera communities and especially the trait composition. Our findings demonstrate the usefulness of considering both taxonomic and functional comp on ents (functio nal diversity) in biodiversity research and we suggest a stronger involvement of plant quality measures in Orthoptera studies.     

Species Richness, Community Specialization and Soil-Vegetation Relationships of Managed Grasslands in a Geologically Heterogeneous Landscape

The increasing importance of the conservation value of managed grasslands has led to many studies exploring edaphic determinants of grassland biodiversity. Most studies, however, come either from very large areas, where biogeographical factors such as dispersal limitation may play a role, or from small, but ecologically rather uniform, regions. In addition, few studies further distinguish between plant specialists and generalists in the interpretation of the observed patterns. Here we studied species richness in semi-natural, managed grasslands in the Strá?ovské vrchy Mountains in the West Carpathians, Slovakia, where there is a matrix of different bedrocks (crystalline, sandstone, claystone, limestone) on a steep altitudinal gradient. In 89 vegetation plots we sampled the species composition of vascular plants and bryophytes and measured soil chemistry, slope angle, heat index, altitude and soil depth. We further applied Ellenberg indicator values and classified species into community specialists or generalists based on the analysis of a large phytosociological database. Using cluster analysis, we delimited five vegetation types that clearly differed in response to soil characteristics. Species richness varied between 19 and 64 species per 16?m². The main compositional gradient correlated with measured soil pH and calcium, but species richness was not significantly correlated with these factors. Soil available phosphorus was not associated with species composition as has been found elsewhere, but it did correlate negatively with species richness and the richness of specialists. Overall, species richness was largely driven by the number of specialists in the plot and particular vegetation types differed conspicuously in their number. We further found significant effects of iron, potassium and sodium on species richness, species composition and the representation of specialists and generalists. Our results provide new insights into the determinants of diversity in managed grasslands as well as to the theoretical species pool concept, explaining species richness variation along a pH gradient.