氮磷养分共同添加促进退化典型草原恢复的机制

土壤养分含量降低是我国草原退化的主要原因之一,养分添加是退化草原恢复的有效措施,但过量养分添加会导致物种多样性降低。为了探讨适宜的养分添加量以及养分添加促进退化草原恢复的机制,本研究选择内蒙古典型草原的退化群落,通过氮(N)磷(P)养分共同添加梯度试验,研究了退化典型草原在群落、功能群和物种3个组织水平上对养分添加的响应。结果表明: 在群落水平,养分添加显著促进了退化典型草原生物量,但没有降低物种多样性;群落生物量随养分添加水平表现为饱和曲线响应,在12.0 g N·m^-2、3.8 g P·m^-2水平趋于饱和;物种多样性在低养分添加水平(N<9.6 g·m^-2、P<3.0 g·m^-2)较对照显著增加,在其余养分添加水平未发生显著变化。在功能群水平,随着养分添加量的增加,多年生根茎禾草在群落中优势度增加,生物量和密度均显著提高;一年生植物生物量和密度在高养分水平添加下显著增加,多年生丛生禾草和杂类草无显著变化。在物种水平,6个物种对养分添加响应不同,羊草通过增加种群密度和个体大小显著增加了种群生物量;大针茅、冰草和糙隐子草种群生物量没有显著变化;星毛委陵菜和黄囊苔草分别因为降低个体大小和种群密度减少了种群生物量。养分添加作为草原恢复的措施,可以显著增加退化草原生物量和物种多样性,降低植物群落中退化指示种,增加多年生根茎禾草。     

Ungulate and topographic control of nitrogen: phosphorus stoichiometry in a temperate grassland; soils, plants and mineralization rates

Although the link between the nitrogen (N): phosphorus (P) stoichiometry of biota and availability has received considerable attention in aquatic systems, there has been relatively little effort to compare the elemental composition of biota and supply in terrestrial habitats. In this study, I explored the effects of a prominent topo-edaphic gradient, from dry hilltop to wet slope-base, and native ungulates on N and P of soils, plants, and rates of in situ net mineralization in grasslands of Yellowstone National Park. Nitrogen and P measurements were made May–September, 2000, in paired, grazed and 38–42 year fenced, ungrazed grassland at five topographically variable sites. Similar to findings from other grassland ecosystems, several site factors associated with organic activity, including soil moisture, C, and plant biomass, covaried with soil N concentration and/or net N mineralization. Soil P concentration and net P mineralization, however, were unrelated to those factors. Instead, net P mineralization was negatively related to soil pH, which is known to control the form of inorganic P and its availability, and soil P was uncorrelated with any soil or plant variable measured in the study. Because of being influenced by different soil properties, N and P net mineralization were unrelated among grasslands. Furthermore, supply and plant N:P ratios were uncorrelated in this grassland system. Based on critical N:P ratios reflecting nutritional limitation of plants, Yellowstone grassland vegetation ranged from being N limited to N-P co-limited. Grazers increased N-P co-limitation by enhancing plant N concentrations and the soil pH gradient across grassland sites regulated plant nutritional limitation by affecting plant-available P. These findings showed how ungulates and a landscape factor, i.e. soil pH, determined plant nutrient status among YNP grasslands differently by influencing plant N concentration versus plant P concentration, respectively.     

Nutrient concentrations and fibre contents of plant community biomass reflect species richness patterns along a broad range of land-use intensities among agricultural grasslands

Understanding changes in biodiversity in agricultural landscapes in relation to land-use type and intensity is a major issue in current ecological research. In this context nutrient enrichment has been identified as a key mechanism inducing species loss in Central European grassland ecosystems. At the same time, insights into the linkage between agricultural land use and plant nutrient status are largely missing. So far, studies on the relationship between chemical composition of plant community biomass and biodiversity have mainly been restricted to wetlands and all these studies neglected the effects of land use. Therefore, we analyzed aboveground biomass of 145 grassland plots covering a gradient of land-use intensities in three regions across Germany. In particular, we explored relationships between vascular plant species richness and nutrient concentrations as well as fibre contents (neutral and acid detergent fibre and lignin) in the aboveground community biomass.We found the concentrations of several nutrients in the biomass to be closely linked to plant species richness and land use. Whereas phosphorus concentrations increased with land-use intensity and decreased with plant species richness, nitrogen and potassium concentrations showed less clear patterns. Fibre fractions were negatively related to nutrient concentrations in biomass, but hardly to land-use measures and species richness. Only high lignin contents were positively associated with species richness of grasslands. The N:P ratio was strongly positively related to species richness and even more so to the number of endangered plant species, indicating a higher persistence of endangered species under P (co-)limited conditions. Therefore, we stress the importance of low P supply for species-rich grasslands and suggest the N:P ratio in community biomass to be a useful proxy of the conservation value of agriculturally used grasslands.     

Characterisation of the type and extent of nutrient limitation in grassland vegetation using a bioassay with intact sods

An important methodological problem in plant ecology concerns the way in which the type and extent of nutrient limitation in terrestrial communities should be assessed. Conclusions on nutrient limitation have been founded mainly on soil extractions, fertiliser trials and tissue nutrient concentrations. In order to avoid some of the problems associated with these methods, we employed a special technique using intact sods which rooted both in the intact soil and in a nutrient solution, from which N, P and K were omitted stepwise. The method was applied to hay-field communities which differed in their history of fertiliser application. Four fields were compared which were not fertilised for 2, 6, 19 or 45 years, while hay making continued. This was done to restore former species-rich grassland communities. We tested the hypothesis that the increase in species diversity in these grasslands was attended by an increase in the number of limiting nutrients.We observed clear shifts in the type and extent of nutrient limitation. Fields which were recently fertilised were characterised by nitrogen and potassium limitation, while phosphorus limitation increased in importance towards the later stages of succession. In the last field (45 years unfertilised) N, P and K equally limited production at the community level. These conclusions differed from those drawn from a fertiliser trial in these same four fields, which failed to detect phosphorus limitation.It is concluded that the use of this method provides a valuable extra source of information while studying relationships between nutrient limitation and species diversity in grassland communities.     

Nitrogen, phosphorus and potassium nutritional status of semiarid steppe grassland in Inner Mongolia

In grazed semiarid steppe ecosystems, much attention has been paid to aspects of growth limitation by water. So far, potential limitation of primary production by plant nutrients was rarely considered. This knowledge is essential for identification of sustainable land-use practices in these large and important ecosystems on the background of over-exploitation and climate change. In the present study plant nutrient concentrations and ratios were investigated with factorial additions of water and N fertilizer at two sites with contrasting soil nutrient availability. Combined analysis of nutrient concentrations, contents, biomass production, and plant N:P ratios consistently confirmed primary growth limitation by water and a strong N limitation when sufficient amounts of water were supplied. P limitation only occurred at the site with low P availability when in addition to the natural supply, water and N fertilizer were given. According to reported thresholds of N:K and K:P ratios, K was not limiting in any plot. The observed nutritional patterns in the plant community were related to the dynamics of species composition and their specific nutrient status. Stipa grandis had the highest N:P ratio whereas Artemisia frigida showed lowest N:P. These nutrient characteristics were related to growth strategies of dominant species. Accordingly, the relative biomass contribution of S. grandis and A. frigida strongly affected the nutrient status of the plant community. Plant N:P ratios indicate the relative limitation by N or P in the semiarid grasslands under sufficient water supply, but other methods of nutritional diagnosis should be used when plant N:P ratios remain below critical values.     

Nutrient limitation in species‐rich Calthion grasslands in relation to opportunities for restoration in a peat meadow landscape

Questions: Which nutrient(s) limit(s) vegetation productivity in Calthion grasslands? Is phosphorus release a bottleneck for restoration of species‐rich Calthion grasslands on rewetted dairy meadows? Location: Three species‐rich Calthion grasslands in the Western Peat District in the Netherlands. Methods: We conducted a field fertilization experiment with nitrogen (N), phosphorus (P) and potassium (K) in three existing Calthion grasslands to evaluate the potential for restoration on rewetted dairy meadows. Responses of above‐ground biomass, tissue nutrient concentrations and nutrient ratios were determined after 2 yr of fertilization. Results: Biomass increased with fertilization with N‐only and K‐only but did not react to P‐only additions. Comparisons of tissue nutrient concentrations and nutrient ratios also gave indications of N and K limitation. Conclusions: The strong P release expected after rewetting should not necessarily interfere with restoration of Calthion communities on rewetted dairy meadows. It is concluded that for successful restoration management measures should focus on reducing N and/or K availability. Potassium might be an overlooked bottleneck in the restoration of species‐rich grasslands.     

Variation in nitrogen and phosphorus concentrations of wetland plants

The use of nutrient concentrations in plant biomass as easily measured indicators of nutrient availability and limitation has been the subject of a controversial debate. In particular, it has been questioned whether nutrient concentrations are mainly species' traits or mainly determined by nutrient availability, and whether plant species have similar or different relative nutrient requirements. This review examines how nitrogen and phosphorus concentration and the N:P ratio in wetland plants vary among species and sites, and how they are related to nutrient availability and limitation. We analyse data from field studies in European non-forested wetlands, from fertilisation experiments in these communities and from growth experiments with wetland plants. Overall, the P concentration was more variable than the N concentration, while variation in N:P ratios was intermediate. Field data showed that the N concentration varies more among species than among sites, whereas the N:P ratio varies more among sites than among species, and the P concentration varies similarly among both. Similar patterns of variation were found in fertilisation experiments and in growth experiments under controlled nutrient supply. Nutrient concentrations and N:P ratios in the vegetation were poorly correlated with various measures of nutrient availability in soil, but they clearly responded to fertilisation in the field and to nutrient supply in growth experiments. In these experiments, biomass N:P ratios ranged from 3 to 40 and primarily reflected the relative availabilities of N and P, although N:P ratios of plants grown at the same nutrient supply could vary three-fold among species. The effects of fertilisation with N or P on the biomass production of wetland vegetation were well related to the N:P ratios of the vegetation in unfertilised plots, but not to N or P concentrations, which supports the idea that N:P ratios, rather than N or P concentrations, indicate the type of nutrient limitation. However, other limiting or stressing factors may influence N:P ratios, and the responses of individual plant species to fertilisation cannot be predicted from their N:P ratios. Therefore, N:P ratios should only be used to assess which nutrient limits the biomass production at the vegetation level and only when factors other than N or P are unlikely to be limiting.     

Grazing-induced changes in plant species composition affect plant and soil properties of grassland mesocosms

Grazing-induced floristic changes in plant communities may accelerate or reduce plant and soil processes through changes in litter quality. Here, we intended to elucidate if the joint action of live and senescing plant tissue of palatable and non-palatable species differentially influences soil processes and properties. We conducted a 1-year experiment with mesocosms from a subhumid grassland. Mesocosms were monocultures of palatable or non-palatable species and a multispecific control. Palatable species included a legume and annual and perennial grasses, whereas non-palatable species included a perennial grass and annual and perennial forbs. Palatable monocultures showed greater soil mineral nitrogen, soil bacterial diversity, and lower soil pH than non-palatable monocultures. These differences were not accounted for by differences in plant biomass. The multispecific control treatment only exhibited greater shoot biomass than the monocultures, and lower root biomass than the palatable monocultures. Our results suggest that the whole (live + dead plant tissue) had a specific imprint on soil system even when variation was not very apparent in terms of plant biomass, and that this effect was associated with plant palatability to domestic large herbivores.     

Plant biomass production and soil nitrogen in mixtures and monocultures of old field Mediterranean annuals

We examine the relationship between plant diversity and ecosystem properties in a Mediterranean grassland. Five legumes, three grasses and two forb species are grown in monocultures and compared with mixtures that include these ten species. Trifolium angustifolium L. (a legume), Lolium rigidum Gaudin (a grass), and Centaurea solstitialis L. (a forb), are replicated in monocultures. Plant cover, root length and biomass, and concentrations of soil nitrate and ammonium are measured in all plots in March and May. Aboveground biomass is measured at a final harvest in late May to early June. Root biomass is significantly higher in the species mixtures than the average of the monocultures. Plant cover and root length are marginally significantly higher (0.05 < P ≤ 0.1) in the mixtures compared to the average of the monocultures. Soil inorganic nitrogen concentrations and aboveground biomass do not significantly differ between the average of the monocultures and the mixtures. Aboveground biomass in T. angustifolium monocultures is significantly higher than in the mixtures, and on average the legume monocultures do not differ significantly from the mixtures. Root length and biomass in L. rigidum monocultures are higher than in the mixtures in March. Nitrate concentrations (which are negatively correlated with root length and biomass) are the lowest in C. solstitialis in May. Thus, we have evidence that some of the measures of ecosystem performance decline in the average of the monocultures when compared with the mixtures, but mixtures never outperform or do more poorly than the best performing monocultures.     

Enhancement of late successional plants on ex-arable land by soil inoculations

Restoration of species-rich grasslands on ex-arable land can help the conservation of biodiversity but faces three big challenges: absence of target plant propagules, high residual soil fertility and restoration of soil communities. Seed additions and top soil removal can solve some of these constraints, but restoring beneficial biotic soil conditions remains a challenge. Here we test the hypotheses that inoculation of soil from late secondary succession grasslands in arable receptor soil enhances performance of late successional plants, especially after top soil removal but pending on the added dose. To test this we grew mixtures of late successional plants in arable top (organic) soil or in underlying mineral soil mixed with donor soil in small or large proportions. Donor soils were collected from different grasslands that had been under restoration for 5 to 41 years, or from semi-natural grassland that has not been used intensively. Donor soil addition, especially when collected from older restoration sites, increased plant community biomass without altering its evenness. In contrast, addition of soil from semi-natural grassland promoted plant community evenness, and hence its diversity, but reduced community biomass. Effects of donor soil additions were stronger in mineral than in organic soil and larger with bigger proportions added. The variation in plant community composition was explained best by the abundances of nematodes, ergosterol concentration and soil pH. We show that in controlled conditions inoculation of soil from secondary succession grassland into ex-arable land can strongly promote target plant species, and that the role of soil biota in promoting target plant species is greatest when added after top soil removal. Together our results point out that transplantation of later secondary succession soil can promote grassland restoration on ex-arable land.     

青藏高原高寒草地AM真菌分布及其对近自然恢复的生态作用

超载过牧以及全球气候变化等导致大部分青藏高原高寒草地呈现持续退化态势。青藏高原高寒草地退化致使地上植物群落逐渐发生更替,地下土壤微生物群落多样性和丰富度发生改变。本文旨在探析青藏高原高寒草地丛枝菌根（arbuscular mycorrhizal,AM）真菌的分布特征、对近自然恢复的生理生态效应及其作用机制。青藏高原高寒草地中已报道4目14属61种AM真菌,约占已知AM真菌物种的20%。高寒草地禾本科植物根围AM真菌物种丰度最高,而莎草科植物根围AM真菌孢子密度最高。3种高寒草地植被类型中,高寒草原AM真菌丰度最高（33种）,山地灌丛草原次之（32种）,高寒草甸最低（22种）。高寒草原以光壁无梗囊霉Acaulospora laevis和闪亮和平囊霉Pacispora scintillans为优势种,山地灌丛草原以摩西斗管囊霉Funneliformis mosseae为优势种,高寒草甸以光壁无梗囊霉A. laevis、近明球囊霉Claroideoglomus claroideum和闪亮和平囊霉P. scintillans为优势种。高寒草地土著AM真菌与植物构建的菌根网络可以通过调节营养元素吸收、分配,促进植物建植和生长;但是毒杂草入侵可以改变土著AM真菌物种多样性和菌根网络,限制本地植被的实际生态位扩张。退化高寒草地中,AM真菌群落具有高的环境适应性和恢复力,其不仅调控地上植物群落建植和多样性,同时AM真菌建植也增加了代谢产物-球囊霉素相关土壤蛋白产生,进而协同改善地下土壤微生态系统,为退化高寒草地早期植被恢复塑造土壤生境。因此,AM真菌在退化高寒草地近自然恢复中具有较大的应用潜力。     

Maintenance of leaf N controls the photosynthetic CO2 response of grassland species exposed to 9 years of free‐air CO2 enrichment

Determining underlying physiological patterns governing plant productivity and diversity in grasslands are critical to evaluate species responses to future environmental conditions of elevated CO₂ and nitrogen (N) deposition. In a 9‐year experiment, N was added to monocultures of seven C₃ grassland species exposed to elevated atmospheric CO₂ (560 μmol CO₂ mol^?1) to evaluate how N addition affects CO₂ responsiveness in species of contrasting functional groups. Functional groups differed in their responses to elevated CO₂ and N treatments. Forb species exhibited strong down‐regulation of leaf N_mass concentrations (?26%) and photosynthetic capacity (?28%) in response to elevated CO₂, especially at high N supply, whereas C₃ grasses did not. Hence, achieved photosynthetic performance was markedly enhanced for C₃ grasses (+68%) in elevated CO₂, but not significantly for forbs. Differences in access to soil resources between forbs and grasses may distinguish their responses to elevated CO₂ and N addition. Forbs had lesser root biomass, a lower distribution of biomass to roots, and lower specific root length than grasses. Maintenance of leaf N, possibly through increased root foraging in this nutrient‐poor grassland, was necessary to sustain stimulation of photosynthesis under long‐term elevated CO₂. Dilution of leaf N and associated photosynthetic down‐regulation in forbs under elevated [CO₂], relative to the C₃ grasses, illustrates the potential for shifts in species composition and diversity in grassland ecosystems that have significant forb and grass components.     

Resilience and restoration of tropical and subtropical grasslands,savannas, and grassy woodlands

Despite growing recognition of the conservation values of grassy biomes, our understanding of how to maintain and restore biodiverse tropical grasslands (including savannas and open‐canopy grassy woodlands) remains limited. To incorporate grasslands into large‐scale restoration efforts, we synthesised existing ecological knowledge of tropical grassland resilience and approaches to plant community restoration. Tropical grassland plant communities are resilient to, and often dependent on, the endogenous disturbances with which they evolved – frequent fires and native megafaunal herbivory. In stark contrast, tropical grasslands are extremely vulnerable to human‐caused exogenous disturbances, particularly those that alter soils and destroy belowground biomass (e.g. tillage agriculture, surface mining); tropical grassland restoration after severe soil disturbances is expensive and rarely achieves management targets. Where grasslands have been degraded by altered disturbance regimes (e.g. fire exclusion), exotic plant invasions, or afforestation, restoration efforts can recreate vegetation structure (i.e. historical tree density and herbaceous ground cover), but species‐diverse plant communities, including endemic species, are slow to recover. Complicating plant‐community restoration efforts, many tropical grassland species, particularly those that invest in underground storage organs, are difficult to propagate and re‐establish. To guide restoration decisions, we draw on the old‐growth grassland concept, the novel ecosystem concept, and theory regarding tree cover along resource gradients in savannas to propose a conceptual framework that classifies tropical grasslands into three broad ecosystem states. These states are: (1) old‐growth grasslands (i.e. ancient, biodiverse grassy ecosystems), where management should focus on the maintenance of disturbance regimes; (2) hybrid grasslands, where restoration should emphasise a return towards the old‐growth state; and (3) novel ecosystems, where the magnitude of environmental change (i.e. a shift to an alternative ecosystem state) or the socioecological context preclude a return to historical conditions.     

Increase of fast nutrient cycling in grassland microcosms through insect herbivory depends on plant functional composition and species diversity

Nutrient cycling in terrestrial ecosystems is affected by various factors such as plant diversity and insect herbivory. While several studies suggest insect herbivory to depend on plant diversity, their interacting effect on nutrient cycling is unclear. In a greenhouse experiment with grassland microcosms of one to six plant species of two functional groups (grasses and legumes), we tested the influence of plant species richness (diversity) and functional composition on plant community biomass production, insect foliar herbivory, soil microbial biomass, and nutrient concentrations in throughfall. To manipulate herbivory, zero, three or six generalist grasshoppers (Chorthippus parallelus) were added to the plant communities. Increasing plant species richness increased shoot biomass and grasshopper performance, without significantly affecting root biomass or insect herbivory. Plant functional composition affected all of these parameters, e.g. legume communities showed the highest shoot biomass, the lowest grasshopper performance and suffered the least herbivory. Nutrient concentrations (dissolved mineral N, PO₄‐P, SO₄‐S) and pH in throughfall increased with herbivory. PO₄‐P and pH increases were positively affected by plant diversity, especially under high herbivore pressure. Plant functional composition affected several throughfall variables, sometimes fully explaining diversity effects. Increasing plant diversity tended to increase soil microbial biomass, but only under high herbivore pressure. Faeces quantities strongly correlated with changes in pH and PO₄‐P; frass may therefore be an important driver of throughfall pH and a main source of PO₄‐P released from living plants. Our results indicate that insect herbivory may significantly influence fast nutrient cycling processes in natural communities, particularly so in managed grasslands.     

降水量变化对藏北高寒草地养分限制的影响

生态系统净初级生产养分限制的模式是现代生态学关注的重要问题。养分的可利用性是草原生态系统生产力动态变化的关键决定因素, 但土壤养分可利用性与整个生态系统中养分限制之间的关系尚不清楚。该研究通过在藏北降水梯度上4种类型高寒草地(从东到西依次是高寒草甸、高寒草甸草原、高寒草原和高寒荒漠草原)设置氮磷养分添加试验, 系统研究氮磷养分添加对不同类型高寒草地的影响, 并探讨降水梯度上高寒草地的氮磷限制模式。结果表明: (1)氮磷添加对不同高寒草地的影响存在差异: 氮添加显著提高了高寒草甸和高寒草甸草原地上生产力, 而对高寒草原和高寒荒漠草原无影响; 单独磷添加对4种高寒草地均无显著影响, 而氮磷添加对4种高寒草地地上生产力均有促进作用。(2)通过计算氮磷共同限制指数发现: 随着降水量减少, 高寒草地氮限制指数从1.18逐渐降低到0.52-0.64, 养分限制模式从氮限制过渡到氮磷共同限制; 磷限制指数在高寒草甸草原和高寒草原为负值, 说明单独磷添加对高寒草甸的生产力有负向作用, 高寒草甸主要受氮限制; 高寒草甸草原介于氮限制与氮磷限制之间, 受到氮磷共同限制, 单独磷添加有负向作用; 高寒荒漠草原受到氮磷共同限制。研究表明, 高寒草地氮磷限制模式存在环境梯度上的递变规律, 随着降水量减少, 高寒草地养分限制模式从氮限制逐渐过渡到氮磷共同限制。由此推断, 未来气候变化条件下氮沉降增加对不同类型高寒草地的影响可能存在差异。同时, 利用养分添加恢复不同类型退化高寒草地时也应将氮磷限制模式的差异考虑进去。     

Plant invasion alters nitrogen cycling by modifying the soil nitrifying community

Plant invasions have dramatic aboveground effects on plant community composition, but their belowground effects remain largely uncharacterized. Soil microorganisms directly interact with plants and mediate many nutrient transformations in soil. We hypothesized that belowground changes to the soil microbial community provide a mechanistic link between exotic plant invasion and changes to ecosystem nutrient cycling. To examine this possible link, monocultures and mixtures of exotic and native species were maintained for 4 years in a California grassland. Gross rates of nitrogen (N) mineralization and nitrification were quantified with ¹⁵N pool dilution and soil microbial communities were characterized with DNA‐based methods. Exotic grasses doubled gross nitrification rates, in part by increasing the abundance and changing the composition of ammonia‐oxidizing bacteria in soil. These changes may translate into altered ecosystem N budgets after invasion. Altered soil microbial communities and their resulting effects on ecosystem processes may be an invisible legacy of exotic plant invasions.     

Effects of plant species richness on stand structure and productivity

Aims Aboveground biomass production commonly increases with species richness in plant biodiversity experiments. Little is known about the direct mechanisms that cause this result. We tested if by occupying different heights and depths above and below ground, and by optimizing the vertical distribution of leaf nitrogen, species in mixtures can contribute to increased resource uptake and, thus, increased productivity of the community in comparison with monocultures.Methods We grew 24 grassland plant species, grouped into four nonoverlapping species pools, in monoculture and 3- and 6-species mixture in spatially heterogeneous and uniform soil nutrient conditions. Layered harvests of above- and belowground biomass, as well as leaf nitrogen and light measurements, were taken to assess vertical canopy and root space structure.Important findings The distribution of leaf mass was shifted toward greater heights and light absorption was correspondingly enhanced in mixtures. However, only some mixtures had leaf nitrogen concentration profiles predicted to optimize whole-community carbon gain, whereas in other mixtures species seemed to behave more 'selfish'. Nevertheless, even in these communities, biomass production increased with species richness. The distribution of root biomass below ground did not change from monocultures to three- and six-species mixtures and there was also no indication that mixtures were better than monocultures at extracting heterogeneously as compared to homogeneously distributed soil resources. We conclude that positive biodiversity effect on aboveground biomass production cannot easily be explained by a single or few common mechanisms of differential space use. Rather, it seems that mechanisms vary with the particular set of species combined in a community.     

Adaptation of selected trees and grasses to low availability of phosphorus

High-nutrient-adapted and low-nutrient-adapted species of New Zealand tussock grasses (Chionochloa), barley (Hordeum), and several taiga trees were grown at three rates of phosphorus supply. Low-nutrient-adapted species in each group of species had similar (grasses) or lower (trees) capacities for phosphate absorption, were less efficient in producing biomass (i.e. had higher nutrient concentrations), and grew more slowly than high-nutrient-adapted species. I conclude that the major adaptation to low nutrient availability in each of these comparisons is a slow growth rate that reduces the annual nutrient requirement.     

Nutrient concentration ratios and co-limitation in South African grasslands

*Assessing plant nutrient limitation is a fundamental part of understanding grassland dynamics. The ratio of concentrations of nitrogen (N) and phosphorus (P) in vegetation has been proposed as an index of the relative limitation of biomass production by N and P, but its utility has not been tested well in grasslands. *At five sites in Kruger National Park, South Africa, across soil and precipitation contrasts, N and P were added in a factorial design to grass-dominated plots. *Although the N:P ratio of unfertilized vegetation across all sites (5.8) would have indicated that production was N-limited, aboveground production was consistently co-limited by N and P. Aboveground production was still greater in plots fertilized with N and P than in those fertilized with just N, but the N:P ratio did not exceed standard thresholds for P limitation in N-fertilized vegetation. Comparisons among sites showed little pattern between site N:P ratio and relative responses to N and P. *When combined with results from other grassland fertilization studies, these data suggest that the N:P ratio of grasses has little ability to predict limitation in upland grasslands. Co-limitation between N and P appears to be much more widespread than would be predicted from simple assumptions of vegetative N:P ratios.     

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.