高寒草甸不同草地类型功能群多样性及组成对植物群落生产力的影响

对不同类型草地功能群多样性和组成与植物群落生产力之间的关系进行了探讨。结果表明：(1)在矮嵩草(Kobresia humlis)草甸和金露梅(Potentilla froticosa)灌丛中,豆科植物的作用比较明显,而其他功能群植物的作用较弱。(2)在藏嵩草(Kobresia tibetica)沼泽化草甸和小嵩草(K．pygmaca)草甸中,虽然杂类草、C3植物和莎草科植物功能群的生产力占群落初级生产力的比例较大,但二者在统计上没有显著性差异,这表明群落生产力除受物种多样性的影响外,也受物种本身特征和环境资源的影响,更主要的是受到功能群内物种密度和均匀度的影响,即功能群组成比功能群多样性更能说明对生态系统过程的影响。(3)不同类型草地群落植物功能群盖度与群落初级生产力呈显著的线性相关。(4)不同类型草地群落生产力与功能群内物种数的变化均表现为单峰曲线关系,即功能群内物种数处于中间水平时,群落生产力最高。     

高寒草甸不同草地群落物种多样性与生产力关系研究

生态系统的结构和功能、生物多样性与生产力的关系问题是近年来群落生态学中研究的中心问题,其中,生态系统生产力水平是其功能的重要表现形式,用4种不同草地类型探讨自然群落的物种多样性与生产力关系.结果表明,矮嵩草草甸、小嵩草草甸和金露梅灌丛群落中物种多样性与生产力的关系呈线性增加关系,藏嵩草沼泽化草甸群落中线性增加关系不显著,这表明群落生产力除受物种多样性的影响外,也受物种本身特征和环境资源的影响.不同的环境资源和环境异质性是形成群落结构特征、物种多样性分布格局差异的主要原因之一.     

高寒草地植物物种多样性与功能多样性的关系

物种多样性与功能多样性的关系是生态学当前研究的热点问题之一,不同区域典型生态系统物种多样性和功能多样性的关系研究有利于生物多样性保护理论的全面发展。以青藏高原地区的主要草地生态系统—高寒草甸和高寒草原为研究对象,采用4个物种多样性指数(Patrick丰富度指数、Shannon-Weiner多样性指数、Pielou均匀度指数和Simpson优势度指数)和9个功能多样性指数(FAD功能性状距离指数、MFAD功能性状平均距离指数、基于样地的FDp和基于群落的FDc功能树状图指数、FRic功能体积指数、FEve功能均匀度指数、Rao功能离散度常二次熵指数、FDiv功能离散指数、FDis功能分散指数),分析了高寒草地植物物种多样性、功能多样性关系及其与初级生产力的关系,以期阐明3个科学问题:不同草地类型的高寒草地生态系统植物物种多样性和功能多样性有何差异?高寒草地生态系统的植物物种多样性和功能多样性有何关系?高寒草地生态系统物种多样性、功能多样性对生态系统功能的影响有何异同?研究结果表明:(1)与高寒草原相比,高寒草甸具有更高的物种多样性、功能丰富度和功能离散度;(2)高寒草甸中,Patrick丰富度与功能丰富度指数(FAD、MFAD、FDp、FDc)和功能离散度指数(FDiv)的具有较强的相关性,最优拟合方程分别为幂函数和二次多项式函数;(3)高寒草原中,Patrick丰富度与功能丰富度指数(FAD、MFAD、FDp、FDc、FRic)、Shannon指数和Simpson指数与FEve指数的相关性较强,最优拟合方程为二次多项式函数,Pielou指数与FEve指数的相关性较强,最优拟合方程为指数函数;(4)高寒草甸的初级生产力分别与物种丰富度指数Patrick、功能离散指数FDiv具有较强的相关性;而高寒草原的初级生产力与4个物种多样性指数间均具有较强的相关性,与功能离散指数FDiv具有较强的相关性,最佳拟合方程均为二次多项式函数。研究的总体结论为:物种多样性、功能多样性、二者之间的关系以及二者与生态系统服务功能(以初级生产力为例)之间的关系在高寒草甸和高寒草原群落中表现迥异,因此在研究青藏高原高寒草地的生态功能时,不能仅仅测度传统的物种多样性,还应测度与物种多样性、生态功能密切相关的功能多样性。     

Alpine plant functional group responses to fertiliser addition depend on abiotic regime and community composition

Background and aims

We ask how productivity responses of alpine plant communities to increased nutrient availability can be predicted from abiotic regime and initial functional type composition.

Methods

We compared four Caucasian alpine plant communities (lichen heath, Festuca varia grassland, Geranium-Hedysarum meadow, snow bed community) forming a toposequence and contrasting in productivity and dominance structure for biomass responses to experimental fertilization (N, P, NP, Ca) and irrigation for 4–5?years.

Results

The dominant plants in more productive communities monopolized added N and P, at the expense of their neighbors. In three out of four communities, N and P fertilizations gave greater aboveground biomass increase than N or P fertilization alone, indicating overall co-limitation of N and P, with N being most limiting. Relative biomass increase in NP treatment was negatively related to biomass in control plots across the four communities. Grasses often responded more vigorously to P, but sedges to N alone. Finally, we present one of the rare examples of a forb showing a strong N or NP response.

Conclusion

Our findings will help improve our ability to predict community composition and biomass dynamics in cool ecosystems subject to changing nutrient availability as induced by climate or land-use changes.     

高寒矮嵩草草甸物种多样性与功能多样性对初级生产力的影响

自然生态系统中植物物种多样性是否随功能多样性变化而影响初级生产力的问题存在很大争议。在高寒矮嵩草草甸,对不同放牧强度、不同利用格局和土壤资源水平的4种生境（Ⅰ.畜圈：重度放牧、资源丰富;Ⅱ.牧道：中度放牧、资源适中;Ⅲ.人工草地：轻-中度放牧、资源贫乏;Ⅳ.封育草地：不放牧、资源贫乏）中物种多样性与功能多样性的变化及其对初级生产力的影响进行了研究。结果表明：物种多样性在牧道生境最高,符合中度干扰理论的预测;人工草地物种多样性低但功能多样性却最高;封育草地物种多样性和功能多样性均较高;物种多样性与地上生物量呈正相关,而功能多样性则与地上生物量呈负相关。表明功能多样性不随物种多样性而变化,二者可能在一定程度上是相互独立地作用于生产力的。群落生产力和生物多样性受生境资源、物种特征、放牧历史和放牧强度等因素的综合影响。     

Co-limitation of plant primary productivity by nitrogen and phosphorus in a species-rich wooded meadow on calcareous soils

High small-scale species richness of calcareous grasslands is generally thought to result from evening of species competitive potentials by limited N availability, because of relatively low herb N/P ratios in these communities. However, P mobility is low in alkaline soils as well. We studied soil chemistry and productivity of herb and moss layers in a very diverse calcareous meadow (up to 76 vascular plant species per m²) to test the hypotheses of a co-limitation of herb productivity by both soil N and P availabilities and moss productivity primarily by P availability. The effect of nutrient supply on productivity was investigated using both a natural productivity gradient as well as fertilization experiments. We observed strong positive correlations of soil P availability and total soil N with the above-ground productivity of herb layer. A long-term fertilization experiment demonstrated that P alone and N and P together increased productivity of vascular species, and that the productivity continuously declined after cessation of fertilization with the effect of previous fertilization occasionally visible even 14 years after treatment termination. A short-term fertilization experiment further demonstrated that N and P when supplied alone increase productivity of vascular plants, suggesting that both elements were limiting. Furthermore, there was a significant interaction between N and P on productivity, indicating that simultaneous N and P supply increased productivity more than separate nutrient additions. Moss productivity was negatively associated with vascular plant productivity. In particular, N addition decreased moss productivity, but moss productivity did not decline in P addition treatments. P requirements of mosses were larger than those of vascular plants. Our data indicate co-limitation of herb productivity by both soil N and P in this highly diverse grassland, while limitation of moss productivity mainly by P. We suggest that N and P co-limitations are common in calcareous diverse grasslands, and may partly explain the extreme small-scale species diversity in these communities.     

Nonlinear responses of productivity and diversity of alpine meadow communities to degradation北大核心CSCD

Aims The alpine meadow degradation could have profound effects on the grassland productivity. The aim of our study is to clarify the dynamic response of community productivity and species diversity in the process of alpine meadow degradation. Methods In the Institute of Geographic Sciences and Natural Resources Research, Chinese Academy of Sciences, Northern Tibetan Grassland Ecosystem Research Station (Nagqu station), we conducted stages experiments with multiple degradation levels: control, mild degraded meadow, moderate degraded meadow, severe degraded meadow and serious sandy meadow. Important findings The response of aboveground biomass to alpine meadow degradation showed a linear or nonlinear increased response patterns, but the belowground biomass and total biomass decreased nonlinearly. As observed in measurement of aboveground biomass, Margalef index, Simpson index, Shannon-Wiener index and Pielou evenness index also exhibit a nonlinear increased response to degradation. The results of structural equation models showed that belowground biomass has a positive relationship with soil carbon content (p < 0.05) and volume water content (p < 0.1). However, soil nutrient and soil physical properties had no significant impact on aboveground biomass (p < 0.1). Compared with soil physical properties, soil nutrition is an important factor influencing the diversity index. In our study, the nonlinear responses of productivity and diversity of alpine meadow were described by using the multiple levels of degradation in space. The results suggested that aboveground productivity cannot interpret the degree of degradation of alpine meadow, and by contrast, alpine meadow degradation should be measured by the change of plant functional groups, such as edible grasses and poisonous forbs. © 2018 Editorial Office of Chinese Journal of Plant Ecology. All rights reserved.     

高寒草甸植物群落物种多样性和生产力关系的光竞争研究

通过施肥形成的生产力由低到高的过程中,物种多样性往往降低。总体竞争假说认为对所有资源的竞争作用对多样性的影响随着生产力提高而加剧,导致物种多样性的下降;光竞争假说则认为随着生产力提高,种间竞争从低生产力时的地下竞争转向高生产力时的光竞争,是光竞争导致了物种多样性的下降。为了验证这两种假说,本文通过在甘南玛曲高寒草甸的均匀施肥实验,研究了光竞争对高寒草甸植物群落物种多样性和生产力关系的影响。结果表明：(1)随着施肥梯度的增加,大部分植物的生长速率加快,高度和叶面积增加;(2)随着施肥梯度的增加,植物群落地上总的生物量提高,叶面积指数增加,透光率降低,物种多样性减少;(3)个体大小不对称的光竞争导致了高寒草甸植物群落物种多样性随施肥梯度的增加而减少。     

The consequences of multiple resource shifts on the productivity and composition of alpine tundra communities: inferences from a long-term snow and nutrient manipulation experiment

Background: Gradients in the amounts and duration of snowpack and resulting soil moisture gradients have been associated with different plant communities across alpine landscapes.

Aims: The extent to which snow additions could alter plant community structure, both alone and in combination with nitrogen (N) and phosphorus (P) additions, provided an empirical assessment of the strength of these variables on structuring the plant communities of the alpine tundra at Niwot Ridge, Colorado Front Range.

Methods: A long-term snow fence was used to study vegetation changes in responses to snowpack, both alone and in conjunction with nutrient amendments, in plots established in dry and moist meadow communities in the alpine belt. Species richness, diversity, evenness and dissimilarity were evaluated after 20 years of treatments.

Results: Snow additions, alone, reduced species richness and altered species composition in dry meadow plots, but not in moist meadow; more plant species were found in the snow-impacted areas than in nearby controls. Changes in plant community structure to N and N + P additions were influenced by snow additions. Above-ground plant productivity in plots not naturally affected by snow accumulation was not increased, and the positive responses of plant species to nutrient additions were reduced by snow addition. Plant species showed individualistic responses to changes in snow and nutrients, and indirect evidence suggested that competitive interactions mediated responses. A Permanova analysis demonstrated that community dissimilarity was affected by snow, N, and P additions, but with these responses differing by community type for snow and N. Snow influenced community patterns generated by N, and finally, the communities impacted by N + P were significantly different than those affected by the individual nutrients.

Conclusions: These results show that changes in snow cover over a 20-year interval produce measureable changes in community composition that concurrently influence and are influenced by soil nutrient availability. Dry meadow communities exhibit more sensitivity to increases in snow cover whereas moist meadow communities appear more sensitive to N enrichment. This study shows that the dynamics of multiple limiting resources influence both the productivity and composition of alpine plant communities, with, species, life form, and functional traits mediating these responses.     

草甸和沼泽植物群落功能多样性与生产力的关系

功能多样性-生产力关系研究结果支持质量比假说和多样性假说, 但对于这两种假说的适用条件尚有争议。通过对吉林省西部草甸和沼泽植物群落的地上生物量、2个物种多样性指标(物种丰富度和Shannon-Weaver指数)、7种植物性状的两类功能多样性指标(群落权重均值和Rao二次熵), 以及土壤环境因子进行调查测量, 研究了群落功能多样性与生产力的关系。结果表明: 1)功能多样性与生产力的关系比物种多样性与生产力的关系更为密切; 2)功能群落权重均值解释生产力变异的能力好于Rao二次熵, 即优势物种对群落生产力的影响作用更大; 3)水淹条件影响着功能多样性与生产力的关系, 以群落权重均值为基础的质量比假说适于解释草甸群落功能多样性与生产力的关系, 而以Rao二次熵为基础的多样性假说适于解释有强烈环境筛(水淹)的沼泽群落功能多样性与生产力的关系。     

AM真菌物种多样性:生态功能、影响因素及维持机制

AM真菌物种多样性是土壤生态系统生物多样性的重要组分之一。尽管对AM真菌多样性已有多年研究,但是,已有研究绝大多数仅停留在对AM真菌群落种属解析层面上,对AM真菌物种多样性生态功能及维持机制方面的认识较浅。从生态功能、影响因素及维持机制3个方面系统地综述了近年来AM真菌多样性领域的研究进展。认为AM真菌多样性对植物群落生产力的调控机制及结合理论与实践解析AM真菌多样性维持机制是该领域未来的重点研究方向。     

Relationship between functional diversity and productivity in meadow and marsh plant communities

功能多样性-生产力关系研究结果支持质量比假说和多样性假说, 但对于这两种假说的适用条件尚有争议。通过对吉林省西部草甸和沼泽植物群落的地上生物量、2个物种多样性指标(物种丰富度和Shannon-Weaver指数)、7种植物性状的两类功能多样性指标(群落权重均值和Rao二次熵), 以及土壤环境因子进行调查测量, 研究了群落功能多样性与生产力的关系。结果表明: 1)功能多样性与生产力的关系比物种多样性与生产力的关系更为密切; 2)功能群落权重均值解释生产力变异的能力好于Rao二次熵, 即优势物种对群落生产力的影响作用更大; 3)水淹条件影响着功能多样性与生产力的关系, 以群落权重均值为基础的质量比假说适于解释草甸群落功能多样性与生产力的关系, 而以Rao二次熵为基础的多样性假说适于解释有强烈环境筛(水淹)的沼泽群落功能多样性与生产力的关系。     

Contrasting diversity patterns of breeding Anatidae in the Northern and Southern Hemispheres

For sustaining ecosystem functions and services, environmental conservation strategies increasingly target to maintain the multiple facets of biodiversity, such as functional diversity (FD) and phylogenetic diversity (PD), not just taxonomic diversity (TD). However, spatial mismatches among these components of biodiversity can impose challenges for conservation decisions. Hence, understanding the drivers of biodiversity is critical. Here, we investigated the global distribution patterns of TD, FD, and PD of breeding Anatidae. Using null models, we clarified the relative importance of mechanisms that influence Anatidae community. We also developed random forest models to evaluate the effects of environmental variables on the Anatidae TD, FD, and PD. Our results showed that geographical variation in Anatidae diversity is hemispheric rather than latitudinal. In the species‐rich Northern Hemisphere (NH), the three diversity indices decreased with latitude within the tropical zone of the NH, but increased in the temperate zone reaching a peak at 44.5–70.0°N, where functional and phylogenetic clustering was a predominant feature. In the Southern Hemisphere (SH), Anatidae diversity increased poleward and a tendency to overdispersion was common. In NH, productivity seasonality and temperature in the coldest quarter were the most important variables. Productivity seasonality was also the most influential predictor of SH Anatidae diversity, along with peak productivity. These findings suggested that seasonality and productivity, both consistent with the energy‐diversity hypothesis, interact with the varying histories to shape the contrasting hemispheric patterns of Anatidae diversity. Phylogenetic diversity (PD) and FD underdispersion, widespread across the species‐rich, seasonally productive mid‐to‐high latitudes of the NH, reflects a rapid evolutionary radiation and resorting associated with Pleistocene cycles of glaciation. The SH continents (and southern Asia) are characterized by a widespread tendency toward PD and FD overdispersion, with their generally species‐poor communities comprising proportionately more older lineages in thermally more stable but less predictably productive environments.     

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。     

Plant Functional Diversity Can Be Independent of Species Diversity: Observations Based on the Impact of 4-Yrs of Nitrogen and Phosphorus Additions in an Alpine Meadow

Past studies have widely documented the decrease in species diversity in response to addition of nutrients, however functional diversity is often independent from species diversity. In this study, we conducted a field experiment to examine the effect of nitrogen and phosphorus fertilization ((NH₄)₂ HPO₄) at 0, 15, 30 and 60 g m^-2 yr^-1 (F0, F15, F30 and F60) after 4 years of continuous fertilization on functional diversity and species diversity, and its relationship with productivity in an alpine meadow community on the Tibetan Plateau. To this purpose, three community-weighted mean trait values (specific leaf area, SLA; mature plant height, MPH; and seed size, SS) for 30 common species in each fertilization level were determined; three components of functional diversity (functional richness, FRic; functional evenness, FEve; and Rao’s index of quadratic entropy, FRao) were quantified. Our results showed that: (i) species diversity sharply decreased, but functional diversity remained stable with fertilization; (ii) community-weighted mean traits (SLA and MPH) had a significant increase along the fertilization level; (iii) aboveground biomass was not correlated with functional diversity, but it was significantly correlated with species diversity and MPH. Our results suggest that decreases in species diversity due to fertilization do not result in corresponding changes in functional diversity. Functional identity of species may be more important than functional diversity in influencing aboveground productivity in this alpine meadow community, and our results also support the mass ratio hypothesis; that is, the traits of the dominant species influenced the community biomass production.     

氮添加对宁夏荒漠草原植物初级生产力的影响机制

许多研究探索了与全球变化相关的生态系统功能的变化,但对生态系统功能变化的机制与途径了解较少。初级生产力是生态系统功能的重要组分,但关于氮(N)添加下荒漠草原植物群落初级生产力如何变化以及变化机制尚未明确,N是否通过影响生物多样性来影响荒漠草原初级生产力?为此,本研究在荒漠草原开展了为期4年的N添加控制实验(2018—2021年),试验处理包括对照和4个N添加水平(5、10、20和40 g m^-2 a^-1),研究了N添加对荒漠草原物种多样性、功能多样性、初级生产力及其关系的影响。结果表明：(1)N添加处理(2018—2021年)改变了植物物种多样性及功能多样性,但年际间变化趋势不同。N添加处理第四年(2021年)荒漠草原植物功能多样性(Rao指数)、群落加权平均值-株高、功能均匀度和功能离散度均显著增加,而荒漠草原植物物种丰富度和Shannon-Wiener指数均显著降低。(2)N添加可以通过影响物种丰富度和功能多样性进而间接地促进荒漠草原初级生产力,但群落加权性状值-株高对初级生产力的影响是正效应,而物种丰富度和功能离散度对初级生产力的影响是...     

Co-limitation of plant primary productivity by nitrogen and phosphorus in a species-rich wooded meadow on calcareous soils

High small-scale species richness of calcareous grasslands is generally thought to result from evening of species competitive potentials by limited N availability, because of relatively low herb N/P ratios in these communities. However, P mobility is low in alkaline soils as well. We studied soil chemistry and productivity of herb and moss layers in a very diverse calcareous meadow (up to 76 vascular plant species per m²) to test the hypotheses of a co-limitation of herb productivity by both soil N and P availabilities and moss productivity primarily by P availability. The effect of nutrient supply on productivity was investigated using both a natural productivity gradient as well as fertilization experiments. We observed strong positive correlations of soil P availability and total soil N with the above-ground productivity of herb layer. A long-term fertilization experiment demonstrated that P alone and N and P together increased productivity of vascular species, and that the productivity continuously declined after cessation of fertilization with the effect of previous fertilization occasionally visible even 14 years after treatment termination. A short-term fertilization experiment further demonstrated that N and P when supplied alone increase productivity of vascular plants, suggesting that both elements were limiting. Furthermore, there was a significant interaction between N and P on productivity, indicating that simultaneous N and P supply increased productivity more than separate nutrient additions. Moss productivity was negatively associated with vascular plant productivity. In particular, N addition decreased moss productivity, but moss productivity did not decline in P addition treatments. P requirements of mosses were larger than those of vascular plants. Our data indicate co-limitation of herb productivity by both soil N and P in this highly diverse grassland, while limitation of moss productivity mainly by P. We suggest that N and P co-limitations are common in calcareous diverse grasslands, and may partly explain the extreme small-scale species diversity in these communities.     

Random species loss underestimates dilution effects of host diversity on foliar fungal diseases under fertilization

With increasing attention being paid to the consequences of global biodiversity losses, several recent studies have demonstrated that realistic species losses can have larger impacts than random species losses on community productivity and resilience. However, little is known about the effects of the order in which species are lost on biodiversity–disease relationships. Using a multiyear nitrogen addition and artificial warming experiment in natural assemblages of alpine meadow vegetation on the Qinghai‐Tibetan Plateau, we inferred the sequence of plant species losses under fertilization/warming. Then the sequence of species losses under fertilization/warming was used to simulate the species loss orders (both realistic and random) in an adjacently novel removal experiment manipulating plot‐level plant diversity. We explicitly compared the effect sizes of random versus realistic species losses simulated from fertilization/warming on plant foliar fungal diseases. We found that realistic species losses simulated from fertilization had greater effects than random losses on fungal diseases, and that species identity drove the diversity–disease relationship. Moreover, the plant species most prone to foliar fungal diseases were also the least vulnerable to extinction under fertilization, demonstrating the importance of protecting low competence species (the ability to maintain and transmit fungal infections was low) to impede the spread of infectious disease. In contrast, there was no difference between random and realistic species loss scenarios simulated from experimental warming (or the combination of warming and fertilization) on the diversity–disease relationship, indicating that the functional consequences of species losses may vary under different drivers.     

Does biodiversity increase spatial stability in plant community biomass?

We tested the hypothesis that biodiversity decreases the spatial variability of biomass production between subplots taken within experimental grassland plots. Our findings supported this hypothesis if functional diversity (weighted Rao's Q ) was considered. Further analyses revealed that diversity in rooting depth and clonal growth form were the most important components of functional diversity stabilizing productivity. Using species or functional group richness as diversity measures there was no significant effect on spatial variability of biomass production, demonstrating the importance of the biodiversity component considered. Moreover, we found a significant increase in spatial variability of productivity with decreasing size of harvested area, suggesting small-scale heterogeneity as an important driver. The ability of diverse communities to stabilize biomass production across spatial heterogeneity may be due to complementary use of spatial niches. Nevertheless, the positive effect of functional diversity on spatial stability appears to be less pronounced than previously reported effects on temporal stability.     

Different inter‐annual responses to availability and form of nitrogen explain species coexistence in an alpine meadow community after release from grazing

Plant species and functional groups in nitrogen (N) limited communities may coexist through strong eco‐physiological niche differentiation, leading to idiosyncratic responses to multiple nutrition and disturbance regimes. Very little is known about how such responses depend on the availability of N in different chemical forms. Here we hypothesize that idiosyncratic year‐to‐year responses of plant functional groups to availability and form of nitrogen explain species coexistence in an alpine meadow community after release from grazing. We conducted a 6 year N addition experiment in an alpine meadow on the Tibetan Plateau released from grazing by livestock. The experimental design featured three N forms (ammonium, nitrate, and ammonium nitrate), crossed with three levels of N supply rates (0.375, 1.500 and 7.500 g N m^?2 yr^?1), with unfertilized treatments without and with light grazing as controls. All treatments showed increasing productivity and decreasing species richness after cessation of grazing and these responses were stronger at higher N rates. Although N forms did not affect aboveground biomas s at community level, different functional groups did show different responses to N chemical form and supply rate and these responses varied from year to year. In support of our hypothesis, these idiosyncratic responses seemed to enable a substantial diversity and biomass of sedges, forbs, and legumes to still coexist with the increasingly productive grasses in the absence of grazing, at least at low and intermediate N availability regimes. This study provides direct field‐based evidence in support of the hypothesis that idiosyncratic and annually varying responses to both N quantity and quality may be a key driver of community structure and species coexistence. This finding has important implications for the diversity and functioning of other ecosystems with spatial and temporal variation in available N quantity and quality as related to changing atmospheric N deposition, land‐use, and climate‐induced soil warming.