Density may alter diversity–productivity relationships in experimental plant communities

Contemporary biodiversity experiments, in which plant species richness is manipulated and aboveground productivity of the system measured, generally demonstrate that lowering plant species richness reduces productivity. However, we propose that community density may in part compensate for this reduction of productivity at low diversity. We conducted a factorial experiment in which plant functional group richness was held constant at three, while plant species richness increased from three to six to 12 species and community density from 440 to 1050 to 2525 seedlings m⁻². Response variables included density, evenness and above- and belowground biomass at harvest. The density gradient converged slightly during the course of the experiment due to about 10% mortality at the highest sowing density. Evenness measured in terms of aboveground biomass at harvest significantly declined with density, but the effect was weak. Overall, aboveground, belowground and total biomass increased significantly with species richness and community density. However, a significant interaction between species richness and community density occurred for both total and aboveground biomass, indicating that the diversity–productivity relationship was flatter at higher than at lower density. Thus, high species richness enabled low-density communities to reach productivity levels otherwise seen only at high density. The relative contributions of the three functional groups C₃, C₄ and nitrogen-fixers to aboveground biomass were less influenced by community density at high than at low species richness. We interpret the interaction effects between community density and species richness on community biomass by expanding findings about constant yield and size variation from monocultures to plant mixtures.     

The interrelationship between productivity,plant species richness and livestock diet: a question of scale?

Question: What relationship exists between productivity, plant species richness and livestock diet? Are the results dependent on scale? Location: A sheep‐grazed Koelerio‐Corynephoretea sandy habitat of the northern upper Rhine (Germany) as a low productivity model system. Methods: The investigation was carried out for three years at a fine scale (2 m²) and for two years at a broad scale (79 m²). Productivity was measured by means of weighed above‐ground phytomass for fine scale and colour‐infrared (CIR) aerial photographs of the same system for fine and broad scales. For both scales, total numbers of vascular plant species and numbers of endangered vascular plant species were extracted from current vegetation relevés. Additionally, we obtained data on livestock diet (grazed phytomass, crude protein content). Results: Statistical analyses show an influence of the year on all variables; relationships between variables are not significant in every year. Species richness and number of endangered species are negatively related to productivity at fine scale while crude protein content and grazed phytomass are positively related to productivity. At the broad scale the diversity‐productivity relationship shows a ‘hump’ with highest species numbers in middle pioneer stages; numbers of endangered species are highest in all pioneer stages. Conclusions: We found a strong impact of scale and year on the diversity‐productivity relationship. It is inappropriate to analyse only small plots (2 m²), and it is necessary to study different years. This vegetation complex is dependent on grazing impact; thus there is an inversely proportional relationship between nature conservation value (high diversity) and livestock nutrition.     

氮素添加对青藏高原高寒草甸植物群落物种丰富度及其与地上生产力关系的影响

植物种群对有限资源的竞争是决定植物群落物种组成、多样性和生产力等群落结构和功能的主要因素。该文以青藏高原高寒草甸为研究对象, 研究了短期内不同水平的氮素添加对高寒草甸植物群落的影响。结果表明: 1)氮素添加提高了土壤中NO₃^--N等可利用资源的含量, 增加了植物群落植被的盖度, 减小了植被的透光率, 随着施氮量的增加, 群落中物种丰富度显著降低(p < 0.001); 2)氮素添加显著改变了植物群落的地上生产力(p < 0.05), 随着施氮量的增加, 地上生产力呈先增加后降低的变化趋势, 各功能群中禾草生物量显著增加, 而杂类草和豆科植物生物量随施氮量的增加逐渐减少; 3)物种多样性与植被透光率呈线性正相关(p < 0.05); 地上生产力与土壤NO₃^--N含量呈线性正相关(p < 0.05); 物种丰富度与地上生产力之间呈负相关关系。这说明短期内氮素添加通过改变土壤中NO₃^--N等可利用资源的含量而对植物群落物种组成和地上生产力产生影响。     

Effects of grazing exclusion on plant species richness and phytomass accumulation vary across a regional productivity gradient

Question: Does long‐term grazing exclusion affect plant species diversity? And does this effect vary with long‐term phytomass accumulation across a regional productivity gradient? Location: Lowland grassy ecosystems across the state of Victoria, southeast Australia. Methods: Floristic surveys and phytomass sampling were conducted across a broad‐scale productivity gradient in grazing exclusion plots and adjacent grazed areas. Differences in species richness, evenness and life‐form evenness between grazed and ungrazed areas were analysed. The environmental drivers of long‐term phytomass accumulation were assessed using multiple linear regression analysis. Results: Species richness declined in the absence of grazing only at the high productivity sites (i.e. when phytomass accumulation was >500 g m^?2). Species evenness and life‐form evenness also showed a negative relationship with increasing phytomass accumulation. Phytomass accumulation was positively associated with both soil nitrogen and rainfall, and negatively associated with tree cover. Conclusions: Competitive dominance is a key factor regulating plant diversity in productive grassy ecosystems, but canopy disturbance is not likely to be necessary to maintain diversity in less productive systems. The results support the predictions of models of the effects of grazing on plant diversity, such as the dynamic equilibrium model, whereby the effects of herbivory are context‐dependent and vary according to gradients of rainfall, soil fertility and tree cover.     

Effects of resource additions on species richness and ANPP in an alpine meadow community

Aims Theories based on resource additions indicate that plant species richness is mainly determined by the number of limiting resources. However, the individual effects of various limiting resources on species richness and aboveground net primary productivity (ANPP) are less well understood. Here, we analyzed potential linkages between additions of limiting resources, species loss and ANPP increase and further explored the underlying mechanisms.Methods Resources (N, P, K and water) were added in a completely randomized block design to alpine meadow plots in the Qinghai-Tibetan Plateau. Plant aboveground biomass, species composition, mean plant height and light availability were measured in each plot. Regression and analysis of variance were used to analyze the responses of these measures to the different resource-addition treatments.Important findings Species richness decreased with increasing number of added limiting resources, suggesting that plant diversity was apparently determined by the number of limiting resources. Nitrogen was the most important limiting resource affecting species richness, whereas P and K alone had negligible effects. The largest reduction in species richness occurred when all three elements were added in combination. Water played a different role compared with the other limiting resources. Species richness increased when water was added to the treatments with N and P or with N, P and K. The decreases in species richness after resource additions were paralleled by increases in ANPP and decreases in light penetration into the plant canopy, suggesting that increased light competition was responsible for the negative effects of resource additions on plant species richness.     

Functional dominance rather than taxonomic diversity and functional diversity mainly affects community aboveground biomass in the Inner Mongolia grassland

The relationship between biodiversity and productivity has been a hot topic in ecology. However, the relative importance of taxonomic diversity and functional characteristics (including functional dominance and functional diversity) in maintaining community productivity and the underlying mechanisms (including selection and complementarity effects) of the relationship between diversity and community productivity have been widely controversial. In this study, 194 sites were surveyed in five grassland types along a precipitation gradient in the Inner Mongolia grassland of China. The relationships between taxonomic diversity (species richness and the Shannon–Weaver index), functional dominance (the community‐weighted mean of four plant traits), functional diversity (Rao's quadratic entropy), and community aboveground biomass were analyzed. The results showed that (1) taxonomic diversity, functional dominance, functional diversity, and community aboveground biomass all increased from low to high precipitation grassland types; (2) there were significant positive linear relationships between taxonomic diversity, functional dominance, functional diversity, and community aboveground biomass; (3) the effect of functional characteristics on community aboveground biomass is greater than that of taxonomic diversity; and (4) community aboveground biomass depends on the community‐weighted mean plant height, which explained 57.1% of the variation in the community aboveground biomass. Our results suggested that functional dominance rather than taxonomic diversity and functional diversity mainly determines community productivity and that the selection effect plays a dominant role in maintaining the relationship between biodiversity and community productivity in the Inner Mongolia grassland.     

Species identity and composition effects on community productivity in a subtropical forest

Biodiversity experiments have shown that high species richness increases productivity in forests. However, forests are diverse regarding community composition. It is unclear how species identity and species composition could contribute to the variation of productivity and how their effects interact with species richness. Here we used data from a 9-year-old planted forest experiment with 40 species to test the effects of species identity and species composition on productivity. The plots in the experiment were designed with various compositions across richness levels. We found that for the top 25 species compositions with the highest volume increment there were six monocultures (24%), eight 2-species mixtures (32%) and eleven 4-species mixtures (44%). Schima superba monoculture had the highest volume in the last study year and highest volume increment over five years, while 2-species and 4-species mixtures with Nyssa sinensis or Liquidambar formosana resulted in high average above-ground volume accumulation. Most of the high-productivity species were deciduous with arbuscular mycorrhizal (AM) associations. In addition, the percentage of AM species in plots had a positive significant effect on both above-ground volume in 2018 and increment from 2013 to 2018. While the percentage of evergreen species showed no significant effect on above-ground volume increment, it did significantly decrease the above-ground volume in 2018. Overall, our results encourage reforestation management to consider species identity and species compositions of deciduous and/or AM species, such as Schima superba, Nyssa sinensis or Liquidambar formosana in the studied subtropical area, if the purpose is to enhance ecosystem productivity.     

Impact of Land-Use Intensity and Productivity on Bryophyte Diversity in Agricultural Grasslands

While bryophytes greatly contribute to plant diversity of semi-natural grasslands, little is known about the relationships between land-use intensity, productivity, and bryophyte diversity in these habitats. We recorded vascular plant and bryophyte vegetation in 85 agricultural used grasslands in two regions in northern and central Germany and gathered information on land-use intensity. To assess grassland productivity, we harvested aboveground vascular plant biomass and analyzed nutrient concentrations of N, P, K, Ca and Mg. Further we calculated mean Ellenberg indicator values of vascular plant vegetation. We tested for effects of land-use intensity and productivity on total bryophyte species richness and on the species richness of acrocarpous (small & erect) and pleurocarpous (creeping, including liverworts) growth forms separately. Bryophyte species were found in almost all studied grasslands, but species richness differed considerably between study regions in northern Germany (2.8 species per 16 m²) and central Germany (6.4 species per 16 m²) due environmental differences as well as land-use history. Increased fertilizer application, coinciding with high mowing frequency, reduced bryophyte species richness significantly. Accordingly, productivity estimates such as plant biomass and nitrogen concentration were strongly negatively related to bryophyte species richness, although productivity decreased only pleurocarpous species. Ellenberg indicator values for nutrients proved to be useful indicators of species richness and productivity. In conclusion, bryophyte composition was strongly dependent on productivity, with smaller bryophytes that were likely negatively affected by greater competition for light. Intensive land-use, however, can also indirectly decrease bryophyte species richness by promoting grassland productivity. Thus, increasing productivity is likely to cause a loss of bryophyte species and a decrease in species diversity.     

Can root trait diversity explain complementarity effects in a grassland biodiversity experiment?

Aims The positive relationship between plant biodiversity and community productivity is well established. However, our knowledge about the mechanisms underlying these positive biodiversity effects is still limited. One of the main hypotheses is that complementarity in resource uptake is responsible for the positive biodiversity effects: plant species differ in resource uptake strategy, which results in a more complete exploitation of the available resources in space and time when plant species are growing together. Recent studies suggest that functional diversity of the community, i.e. the diversity in functional characteristics ('traits') among species, rather than species richness per se, is important for positive biodiversity effects. However, experimental evidence for specific trait combinations underlying resource complementarity is scarce. As the root system is responsible for the uptake of nutrients and water, we hypothesize that diversity in root traits may underlie complementary resource use and contribute to the biodiversity effects.Methods In a common garden experiment, 16 grassland species were grown in monoculture, 4-species mixtures differing in root trait diversity and 16-species mixtures. The 4-species mixtures were designed to cover a gradient in average rooting depth. Above-ground biomass was cut after one growing season and used as a proxy for plant productivity to calculate biodiversity effects.Important findings Overall, plant mixtures showed a significant increase in biomass and complementarity effects, but this varied greatly between communities. However, diversity in root traits (measured in a separate greenhouse experiment and based on literature) could not explain this variation in complementarity effects. Instead, complementarity effects were strongly affected by the presence and competitive interactions of two particular species. The large variation in complementarity effects and significant effect of two species emphasizes the importance of community composition for positive biodiversity effects. Future research should focus on identifying the traits associated with the key role of particular species for complementarity effects. This may increase our understanding of the links between functional trait composition and biodiversity effects as well as the relative importance of resource complementarity and other underlying mechanisms for the positive biodiversity effects.     

亚热带森林树种多样性对凋落叶分解胞外酶活性的影响

森林生物多样性与生态系统功能关系是当前群落生态学的热点研究领域。然而, 以往研究更多聚焦在森林植物多样性丧失对群落生产力的影响, 而对森林凋落物分解的相关研究稍显不足。森林凋落叶分解的快慢直接受控于凋落物分解者分泌的胞外酶的活性, 后者更是指示森林生态系统养分循环的重要指标之一。本研究依托我国江西亚热带森林生物多样性与生态系统功能控制实验, 通过对不同植物多样性梯度样方内目标树种凋落叶胞外酶活性、理化性质以及腐生真菌的分析, 探索树种多样性丧失对胞外酶活性的影响及其调控机制, 以探讨森林树种多样性对地表、地下生态过程和功能的影响。结果表明, 样方水平树种多样性丧失显著影响胞外酶的活性, 除单种样方外, 随着样方水平树种丰富度的增加, 胞外酶活性呈现出增长趋势; 与碳周转相关的α-葡萄糖苷酶(AG)、β-葡萄糖苷酶(BG)、纤维二糖水解酶(CB)在树种多样性最大时活性达到最高; 而木糖苷酶(XS)以及与氮、磷和顽拗有机养分分解相关的N-乙酰-β-氨基葡萄糖苷酶(NAG)、酸性磷酸酶(AP)和多酚氧化酶(PPO)在树种多样性较低时活性较高。针对目标树种周围的邻居树种多样性进一步分析发现, 各胞外酶活性随着邻居树种多样性的变化呈“单峰”响应趋势, 酶活性大多在邻居树种丰富度为6时呈现峰值。研究发现真菌分解者在胞外酶活性对植物多样性的响应上可能存在重要的调控作用, 可以推测树种多样性通过改变腐生真菌分解者的群落结构和多度, 从而影响胞外酶活性。     

Dominance by an obligate annual affects the morphological characteristics and biomass production of a planted wetland macrophyte community

Aims Biodiversity–ecosystem function experiments can test for causal relationships between planting diversity and community productivity. Planting diversity is routinely introduced as a design element in created wetlands, yet substantive support for the finding that early diversity positively affects ecosystem functioning is lacking for wetlands. We conducted a 2-year diversity–productivity experiment using freshwater wetland mesocosms to investigate community biomass production as affected by planted macrophyte functional richness.Methods A richness gradient of macrophytes in four emergent wetland plant functional groups was established in freshwater mesocosms for two consecutive years. Species-specific aboveground morphological traits of plant size were measured at peak growth in both years; rooting depth was measured for each species in the second year. Aboveground biomass (AGB) and belowground biomass (BGB) were harvested after peak growth in the second year; first year AGB was estimated from morphological traits in constructed regression equations. Net richness effects (i.e. both complementarity effects and selection effects) were calculated using an additive partitioning method.Important findings Species richness had a positive effect on community AGB relative to monocultures in the first year. In the second year, mean AGB was significantly reduced by competition in the most species-rich mixtures and all mixtures underyielded relative to the average monoculture. Competition for soil resources was weaker belowground, whereby root distribution at depths>20cm was reduced at the highest richness levels but overall BGB production was not affected. Changes in species biomass were strongly reflected by variation in species morphological traits, and species above and belowground performances were highly correlated. The obligate annual (Eleocharis obtusa), a dominant competitor, significantly contributed to the depression of perennial species' growth in the second growing season. To foster primary productivity with macrophyte richness in early successional communities of created wetlands where ruderal strategies are favored and competition may be stronger than species complementarity, unsystematic planting designs such as clustering the same or similar species could provide protection for some individuals. Additionally, engineering design elements fostering spatial or temporal environmental variability (e.g. microtopography) in newly created wetlands helps diversify the responses of wetland macrophyte species to their environment and could allow for greater complementarity in biomass production.     

Small mammal herbivores mediate the effects of soil nitrogen and invertebrate herbivores on grassland diversity

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Dispersal and seed limitation affect diversity and productivity of montane grasslands

So far, seed limitation as a local process, and dispersal limitation as a regional process have been largely neglected in biodiversity–ecosystem functioning research. However, these processes can influence both local plant species diversity and ecosystem processes, such as biomass production. We added seeds of 60 species from the regional species pool to grassland communities at 20 montane grassland sites in Germany. In these sites, plant species diversity ranged from 10 to 34 species m⁻² and, before manipulation, diversity was not related to aboveground biomass, which ranged from 108 to 687 g m⁻². One year after seed addition, local plant species richness had increased on average by six species m⁻² (29%) compared with control plots, and this increase was highest in grasslands with intermediate productivity. The increased diversity after adding seeds was associated with an average increase of aboveground biomass of 36 g m⁻² (14.8%) compared with control plots. Thus, our results demonstrate that a positive relationship between changes in species richness and productivity, as previously reported from experimental plant communities, also holds for natural grassland ecosystems. Our results show that local plant communities are dispersal limited and a hump‐shaped model appears to be the limiting outline of the natural diversity–productivity relationship. Hence, the effects of dispersal on local diversity can substantially affect the functioning of natural ecosystems.     

Impacts of nitrogen addition on plant biodiversity in mountain grasslands depend on dose,application duration and climate: a systematic review

Although the influence of nitrogen (N) addition on grassland plant communities has been widely studied, it is still unclear whether observed patterns and underlying mechanisms are constant across biomes. In this systematic review, we use meta‐analysis and metaregression to investigate the influence of N addition (here referring mostly to fertilization) upon the biodiversity of temperate mountain grasslands (including montane, subalpine and alpine zones). Forty‐two studies met our criteria of inclusion, resulting in 134 measures of effect size. The main general responses of mountain grasslands to N addition were increases in phytomass and reductions in plant species richness, as observed in lowland grasslands. More specifically, the analysis reveals that negative effects on species richness were exacerbated by dose (ha^?1 year^?1) and duration of N application (years) in an additive manner. Thus, sustained application of low to moderate levels of N over time had effects similar to short‐term application of high N doses. The climatic context also played an important role: the overall effects of N addition on plant species richness and diversity (Shannon index) were less pronounced in mountain grasslands experiencing cool rather than warm summers. Furthermore, the relative negative effect of N addition on species richness was more pronounced in managed communities and was strongly negatively related to N‐induced increases in phytomass, that is the greater the phytomass response to N addition, the greater the decline in richness. Altogether, this review not only establishes that plant biodiversity of mountain grasslands is negatively affected by N addition, but also demonstrates that several local management and abiotic factors interact with N addition to drive plant community changes. This synthesis yields essential information for a more sustainable management of mountain grasslands, emphasizing the importance of preserving and restoring grasslands with both low agricultural N application and limited exposure to N atmospheric deposition.     

Short-term invasibility patterns in burnt and unburnt experimental Mediterranean grassland communities of varying diversities

This paper reports the findings of a short-term natural invasibility field study in constructed Mediterranean herbaceous communities of varying diversities, under a fire treatment. Three components of invasibility, i.e. species richness, density and biomass of invaders, have been monitored in burnt and unburnt experimental plots with resident diversity ranging from monocultures to 18-species mixtures. In general, species richness, density and biomass of invaders decreased significantly with the increase of resident species richness. Furthermore, the density and biomass of invading species were significantly influenced by the species composition of resident communities. Although aboveground biomass, leaf area index, canopy height and percent bare ground of the resident communities explained a significant part of the variation in the success of invading species, these covariates did not fully explain the effects of resident species richness. Fire mainly influenced invasibility via soil nutrient levels. The effect of fire on observed invasibility patterns seems to be less important than the effects of resident species richness. Our results demonstrate the importance of species richness and composition in controlling the initial stages of plant invasions in Mediterranean grasslands but that there was a lack of interaction with the effects of fire disturbance.     

Leaf litter diversity alters microbial activity,microbial abundances,and nutrient cycling in a subtropical forest ecosystem

Human activities affect both tree species composition and diversity in forested ecosystems. This in turn alters the species diversity of plant litter and litter quality, which may have cascading effects on soil microbial communities and their functions for decomposition and nutrient cycling. We tested microbial responses to litter species diversity in a leaf litter decomposition experiment including monocultures, 2-, and 4-species mixtures in the subtropical climate zone of southeastern China. Soil microbial community composition was assessed by lipid analysis, and microbial functions were measured using extracellular enzyme activity and gross rates of nitrogen mineralization. We observed a positive relationship between litter species diversity and abundances of mycorrhizal fungi and actinomycetes. Alternatively, enzyme activities involved in carbon and phosphorus acquisition, and enzyme indices of relative carbon limitation, were higher only in the 4-species mixtures. This suggests that the minimum basal substrate level for enzyme production was reached, or that limitation was higher, at the highest diversity level only. Responses to litter diversity also changed over time, where phosphatase responses to litter diversity were strongest early in decomposition and the indices of carbon limitation relative to other nutrients showed stronger responses later in decomposition. Enzyme activities were related to lipid biomarker data and the mass of litter remaining at the third time point, but relationships between enzyme activity and the mass of litter remaining were not consistent across other time points. We conclude that litter species richness will likely only reduce microbial functions at key intervals of diversity loss while microbial growth is more sensitive to incremental diversity loss, with no clear relationships between them or to ecosystem functions. The observed litter diversity effects on soil microbial biomass and enzyme activity indicate interactions of aboveground and belowground communities, and together with environmental conditions they are important for maintaining ecosystem functions.     

The roles of community biomass and species pools in the regulation of plant diversity

Considerable debate has developed over the importance of community biomass and species pools in the regulation of community diversity. Attempts to explain patterns of plant diversity as a function of community biomass or productivity have been only partially successful and, in general, have explained only a fraction of the observed variation in diversity. At the same time, studies that have focused on the importance of species pools have led some to conclude that diversity is primarily regulated in the short term by the size of the species pool rather than by biotic interactions. In this paper, I explore how community biomass and species pools may work in combination to regulate diversity in herbaceous plant communities. To address this problem, I employ a simple model in which the dynamics of species richness are a function of aboveground community biomass and environmentally controlled gradients in species pools. Model results lead to two main predictions about the role of biomass regulation: (1) Seasonal dynamics of richness will tend to follow a regular oscillation, with richness rising to peak values during the early to middle portion of the growing season and then declining during the latter part of the season. (2) Seasonal dieback of aboveground tissues facilitates the long‐term maintenance of high levels of richness in the community. The persistence of aboveground tissues and accumulation of litter are especially important in limiting the number of species through the suppression of recruitment. Model results also lead to two main predictions about the role of species pools: (1) The height and position of peak richness relative to community biomass will be influenced by the rate at which the species pool increases as available soil resources increase. (2) Variations in nonresource environmental factors (e.g. soil pH or soil salinity) have the potential to regulate species pools in a way that is uncorrelated with aboveground biomass. Under extreme conditions, such nonresource effects can create a unimodal envelope of biomass–richness values. Available evidence from the literature provides partial support for these predictions, though additional data are needed to provide more convincing tests.     

Earthworm and belowground competition effects on plant productivity in a plant diversity gradient

Diversity is one major factor driving plant productivity in temperate grasslands. Although decomposers like earthworms are known to affect plant productivity, interacting effects of plant diversity and earthworms on plant productivity have been neglected in field studies. We investigated in the field the effects of earthworms on plant productivity, their interaction with plant species and functional group richness, and their effects on belowground plant competition. In the framework of the Jena Experiment we determined plant community productivity (in 2004 and 2007) and performance of two phytometer plant species [Centaurea jacea (herb) and Lolium perenne (grass); in 2007 and 2008] in a plant species (from one to 16) and functional group richness gradient (from one to four). We sampled earthworm subplots and subplots with decreased earthworm density and reduced aboveground competition of phytometer plants by removing the shoot biomass of the resident plant community. Earthworms increased total plant community productivity (+11%), legume shoot biomass (+35%) and shoot biomass of the phytometer C. jacea (+21%). Further, phytometer performance decreased, i.e. belowground competition increased, with increasing plant species and functional group richness. Although single plant functional groups benefited from higher earthworm numbers, the effects did not vary with plant species and functional group richness. The present study indicates that earthworms indeed affect the productivity of semi-natural grasslands irrespective of the diversity of the plant community. Belowground competition increased with increasing plant species diversity. However, belowground competition was modified by earthworms as reflected by increased productivity of the phytometer C. jacea. Moreover, particularly legumes benefited from earthworm presence. Considering also previous studies, we suggest that earthworms and legumes form a loose mutualistic relationship affecting essential ecosystem functions in temperate grasslands, in particular decomposition and plant productivity. Further, earthworms likely alter competitive interactions among plants and the structure of plant communities by beneficially affecting certain plant functional groups.     

Shifting grassland plant community structure drives positive interactive effects of warming and diversity on aboveground net primary productivity

Ecosystems worldwide are increasingly impacted by multiple drivers of environmental change, including climate warming and loss of biodiversity. We show, using a long‐term factorial experiment, that plant diversity loss alters the effects of warming on productivity. Aboveground primary productivity was increased by both high plant diversity and warming, and, in concert, warming (≈1.5 °C average above and belowground warming over the growing season) and diversity caused a greater than additive increase in aboveground productivity. The aboveground warming effects increased over time, particularly at higher levels of diversity, perhaps because of warming‐induced increases in legume and C4 bunch grass abundances, and facilitative feedbacks of these species on productivity. Moreover, higher plant diversity was associated with the amelioration of warming‐induced environmental conditions. This led to cooler temperatures, decreased vapor pressure deficit, and increased surface soil moisture in higher diversity communities. Root biomass (0–30 cm) was likewise consistently greater at higher plant diversity and was greater with warming in monocultures and at intermediate diversity, but at high diversity warming had no detectable effect. This may be because warming increased the abundance of legumes, which have lower root : shoot ratios than the other types of plants. In addition, legumes increase soil nitrogen (N) supply, which could make N less limiting to other species and potentially decrease their investment in roots. The negative warming × diversity interaction on root mass led to an overall negative interactive effect of these two global change factors on the sum of above and belowground biomass, and thus likely on total plant carbon stores. In total, plant diversity increased the effect of warming on aboveground net productivity and moderated the effect on root mass. These divergent effects suggest that warming and changes in plant diversity are likely to have both interactive and divergent impacts on various aspects of ecosystem functioning.     

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.