Plant functional traits improve diversity‐based predictions of temporal stability of grassland productivity

Synthesis The temporal stability of plant production is greater in communities with high than low species richness, but stability also may depend on species abundances and growth‐related traits. Annual precipitation varied by greater than a factor of three over 11 years in central Texas, USA leading to large variation in production. Stability was greatest in communities that were not dominated by few species and in which dominant species rooted shallowly, had dense leaves, or responded to the wettest year with a minimal increase in production. Stability may depend as much on species abundances and functional traits as on species richness alone. Aboveground net primary productivity (ANPP) varies in response to temporal fluctuations in weather. Temporal stability of community ANPP may be increased by increasing plant species richness, but stability often varies at a given richness level implying a dependence on abundances and functional properties of member species. We measured stability in ANPP during 11 years in field plots (Texas, USA) in which we varied the richness and relative abundances of perennial grassland species at planting. We sought to identify species abundance patterns and functional traits linked to the acquisition and processing of essential resources that could be used to improve richness‐based predictions of community stability. We postulated that community stability would correlate with abundance‐weighted indices of traits that influence plant responses to environmental variation. Annual precipitation varied by a factor of three leading to large inter‐annual variation in ANPP. Regression functions with planted and realized richness (species with > 1% of community ANPP during the final four years) explained 32% and 25% of the variance in stability, respectively. Regression models that included richness plus the fraction of community ANPP produced by the two most abundant species in combination with abundance‐weighted values of either the fraction of sampled root biomass at 20–45 cm depth, leaf dry matter content (LDMC), or response to greater‐than‐average precipitation of plants grown in monocultures explained 58–69% (planted richness) and 58–64% (realized richness) of the variance in stability. Stability was greatest in communities that were not strongly dominated by only two species and in which plants rooted shallowly, had high values of LDMC, or responded to the wettest year with a minimal increase in ANPP. Our results indicate that the temporal stability of grassland ANPP may depend as much on species abundances and functional traits linked to plant responses to precipitation variability as on species richness alone.     

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.     

Differential roles of species richness versus species asynchrony in regulating community stability along a precipitation gradient

Plant community may provide products and services to humans. However, patterns and drivers of community stability along a precipitation gradient remain unclear. A regional‐scale transect survey was conducted over a 3‐year period from 2013 to 2015, along a precipitation gradient from 275 to 555 mm and spanning 440 km in length from west to east in a temperate semiarid grassland of northern China, a central part of the Eurasian steppe. Our study provided regional‐scale evidence that the community stability increased with increasing precipitation in the semiarid ecosystem. The patterns of community stability along a precipitation gradient were ascribed to community composition and community dynamics, such as species richness and species asynchrony, rather than the abiotic effect of precipitation. Species richness regulated the temporal mean (μ) of aboveground net primary productivity (ANPP), while species asynchrony regulated the temporal standard deviation (σ) of ANPP, which in turn contributed to community stability. Our findings highlight the crucial role of community composition and community dynamics in regulating community stability under climate change.     

Relationship of Productivity to Species Richness in the Xinjiang Temperate Grassland

The relationship between species richness (SR) and aboveground net primary productivity (ANPP) is still a central and debated issue in community ecology. Previous studies have often emphasized the relationship of alpha diversity (number of species identity) to the mean ANPP with respect to the SR-ANPP relationship while neglecting the contribution of beta diversity (dissimilarity in species composition) to the mean ANPP and to the stability of ANPP (coefficient of ANPP: CV of ANPP). In this study, we used alpha and beta diversity, mean ANPP and the CV of ANPP collected from 159 sites and belonging to three vegetation types in the Xinjiang temperate grassland to first examine their trends along climatic factors and among different vegetation types and then test the relationship among alpha (beta) diversity and mean ANPP and the CV of ANPP. Our results showed that in the Xinjiang temperate grasslands, alpha diversity was positively and linearly correlated with MAP but unimodally correlated with MAT. Meanwhile, beta diversity was unimodally correlated with MAP but linearly correlated with MAT. Relative to desert steppe, meadow steppe and typical steppe had the highest alpha and beta diversity, respectively. Except for ANPP exhibiting a quadratic relationship with MAP, no significant relationship was found among ANPP, the CV of ANPP and climatic factors. ANPP and the CV of ANPP also exhibited no apparent patterns in variation among different vegetation types. Our results further showed that mean ANPP was closely associated with alpha diversity. Both linear and unimodal relationships were detected between alpha diversity and mean ANPP, but their particular form was texture-dependent. Meanwhile, the CV of ANPP was positively correlated with beta diversity. Our results indicated that in addition to incorporating alpha diversity and mean ANPP, incorporating beta diversity and the CV of ANPP could expand our understanding of the SR-ANPP relationship.     

Plant community dynamics in a semi-arid ecosystem in relation to nutrient addition following a major disturbance

Effects of annual additions of mineral N and P (100 kg ha^–1) on plant species composition and annual aboveground net primary production (ANPP) were investigated during the first three years following disturbance in a semi-arid ecosystem. Additions of N reduced richness of perennial plant species during years 2 and 3, while P reduced the number of perennial species only in year 3. From year 1 to year 2, annual and biennial species richness declined in all treatments while ANPP of annual species increased greatly. Added N increased ANPP of annual species while it decreased ANPP of most perennial species relative to the unfertilized control treatment. Community similarities were higher for the control and native vegetation than for other pairs of treatments using both species presence and plant production data. Nitrogen additions have retarded but not completely arrested secondary succession in this system.     

西藏高寒草地物种多样性和生产力的环境驱动机制

全球变暖影响物种多样性和生产力及其关系。关于全球和区域的物种多样性与生产力关系已有许多研究,但气候变暖背景下西藏高寒草地物种多样性与生产力的关系及其环境驱动机制研究仍然很少。基于西藏高寒草地实测的35个样点调查数据,利用回归分析、Pearson相关性分析、方差分解和结构方程模型等方法,探究了物种多样性和生产力的关系及其影响机制。研究发现:(1)高寒草地的物种丰富度指数和香农-威纳指数与地上植被净初级生产力(ANPP)呈显著正相关关系,且ANPP对物种丰富度指数的变化更为敏感;(2)物种丰富度指数与经度、土壤有效氮、土壤有效磷、年降水量呈显著正相关;Shannon-Wiener指数与海拔和纬度呈显著负相关,与年均温度呈显著正相关;ANPP与经度、土壤有效氮、年降水量和年均温度呈显著正相关,与海拔、纬度和土壤有效钾呈显著负相关;(3)地理因子、土壤养分和气候因子的交互作用对物种丰富度指数和ANPP的贡献率最大,分别为10.99%和32.91%,地理因子和气候因子的交互作用对Shannon-Wiener指数的贡献率最大,为13.61%;(4)地理因子通过调控土壤养分和气候因子间接影响物种多样性和ANPP,土壤养分和气候因子均直接影响物种多样性和ANPP。研究结果揭示了环境因子对物种多样性和生产力的综合调控机制,为西藏高寒草地生态系统科学应对气候变化提供了依据。     

Fertilization effects on species density and primary productivity in herbaceous plant communities

Fertilization experiments in plant communities are often interpreted in the context of a hump-shaped relationship between species richness and productivity. We analyze results of fertilization experiments from seven terrestrial plant communities representing a productivity gradient (arctic and alpine tundra, two old-field habitats, desert, short- and tall-grass prairie) to determine if the response of species richness to experimentally increased productivity is consistent with the hump-shaped curve. In this analysis, we compared ratios of the mean response in nitrogen-fertilized plots to the mean in control plots for aboveground net primary productivity (ANPP) and species density ( D ; number of species per plot of fixed unit area). In general, ANPP increased and plant species density decreased following nitrogen addition, although considerable variation characterized the magnitude of response. We also analyzed a subset of the data limited to the longest running studies at each site (≥4 yr), and found that adding 9 to 13 g N m⁻² yr⁻¹ (the consistent amount used at all sites) increased ANPP in all communities by approximately 50% over control levels and reduced species density by approximately 30%. The magnitude of response of ANPP and species density to fertilization was independent of initial community productivity. There was as much variation in the magnitude of response among communities within sites as among sites, suggesting community-specific mechanisms of response. Based on these results, we argue that even long-term fertilization experiments are not good predictors of the relationship between species richness and productivity because they are relatively small-scale perturbations whereas the pattern of species richness over natural productivity gradients is influenced by long-term ecological and evolutionary processes.     

How enrichment,ecosystem size,and their effects on species richness co‐determine the stability of microcosm communities

Nutrient enrichment, ecosystem size, and richness each may directly affect the stability of both populations and communities. Alternatively, nutrient enrichment and ecosystem size each may directly affect richness, which in turn may affect stability. No previous studies, however, have tested empirically how these three factors interact and co‐determine stability. We manipulated nutrient input and ecosystem size in replicate microcosms containing a diverse bacterial flora, and a range of green algae and heterotrophic protozoa, and used these manipulations and the resulting variation in species richness to measure their combined effects on temporal stability of both populations and communities. Results showed that nutrient enrichment and ecosystem size controlled protist richness, and their effects on stability could be mediated by richness. In addition, both community‐level and population‐level stability increased with protist richness. Furthermore, mean species evenness and mean species richness was negatively related. Effects of statistical averaging, overyielding, and component population stability were identified as possible mechanisms involved explaini ng the stabilizing effects of richness on community stability. Their relative strength in influencing stability, however, is likely to change as mean evenness decreased with increasing richness. This decrease in evenness would tend to weaken the strength of the statistic averaging effect, but increase the strength of the other two mechanisms due to relatively lower population variability (component population stability) and higher mean biovolumes of dominant protists (overyielding).     

刈割、施肥对高寒草甸物种多样性与生态系统功能关系的影响及群落稳定性机制

植物群落中不同“功能身份”物种的多样性与特定生态系统功能之间具有何种关系及其作用机制尚不明确。通过在高寒矮嵩草(Kobresia humilis)草甸为期5年的刈割(不刈割、留茬3 cm、留茬1 cm)、施肥(施肥、不施肥)和浇水(浇水、不浇水)控制实验, 研究了刈割与土壤资源获得性梯度上不同“功能身份”物种(群落中所有物种、响应物种、作用物种和共有物种)的多样性变化与群落地上净初级生产力和稳定性的关系以及稳定性机制。研究结果显示: 群落中响应物种、作用物种和共有物种数分别占全部物种数的36.6%、18.3%和64.8%, 物种多样性对生态系统功能具有不同的效应, 净初级生产力主要受响应物种和作用物种的多样性变化影响, 而稳定性则主要由共有物种的多样性变化决定; 群落稳定性的维持主要依赖于共有物种的多样性增加, 其作用机制是投资组合效应, 而超产效应和异步性效应对稳定性并无作用; 刈割和施肥对物种多样性、稳定性和净初级生产力具有相反的影响, 前者能增加物种多样性和稳定性, 并降低净初级生产力, 而后者的作用正相反。这与群落中全部物种的多样性变化受刈割影响较大, 而作用物种的多样性变化受资源获得性影响较大有关。上述结果表明高寒草甸生态系统地上净初级生产力主要由少数影响生产力的作用物种的多样性决定, 而稳定性则由大量共有物种的多样性所掌控。投资组合效应是物种多样性导致稳定性的机制。由于群落中不同物种的多样性效应具有分异性, 对于特定的生态系统功能而言, 物种的“功能身份”可能比物种多样性本身更重要, 不加区别地笼统定义物种多样性与生态系统功能的关系可能欠妥。     

Non-neutral community dynamics: empirical predictions for ecosystem function and diversity from linearized consumer–resource interactions

A general model of linearized species interactions, essentially Lotka–Volterra theory, applied to questions of biodiversity has previously been shown to be a powerful tool for understanding local species–abundance patterns and community responses to environmental change for a single trophic level. Here this approach is extended to predict community composition and responses to environmental changes in trophically structured systems. We show how resource and consumer species richness and their relative abundances vary with the means and variances in enrichment level and strengths of intra- and interspecific interactions. Also demonstrated are the responses of local resource and consumer species richness to the global species pools at both trophic levels, as well as the covariation with net resource productivity. These predictions for resource and consumer specific responses to changes in environmental enrichment and global biodiversity are directly testable.     

The potential bias of nitrogen deposition effects on primary productivity and biodiversity

Atmospheric nitrogen (N) deposition is composed of both inorganic nitrogen (IN) and organic nitrogen (ON), and these sources of N may exhibit different impacts on ecosystems. However, our understanding of the impacts of N deposition is largely based on experimental gradients of INs or more rarely ONs. Thus, the effects of N deposition on ecosystem productivity and biodiversity may be biased. We explored the differential impacts of N addition with different IN:ON ratios (0:10, 3:7, 5:5, 7:3, and 10:0) on aboveground net primary productivity (ANPP) of plant community and plant diversity in a typical temperate grassland with a long-term N addition experiment. Soil pH, litter biomass, soil IN concentration, and light penetration were measured to examine the potential mechanisms underlying species loss with N addition. Our results showed that N addition significantly increased plant community ANPP by 68.33%–105.50% and reduced species richness by 16.20%–37.99%. The IN:ON ratios showed no significant effects on plant community ANPP. However, IN-induced species richness loss was about 2.34 times of ON-induced richness loss. Soil pH was positively related to species richness, and they exhibited very similar response patterns to IN:ON ratios. It implies that soil acidification accounts for the different magnitudes of species loss with IN and ON additions. Overall, our study suggests that it might be reasonable to evaluate the effects of N deposition on plant community ANPP with either IN or ON addition. However, the evaluation of N deposition on biodiversity might be overestimated if only IN is added or underestimated if only ON is added.     

Testing the effects of plant species loss on multiple ecosystem functions based on extinction scenarios

Ecosystem functions are threatened by continuing global loss of biodiversity. We simultaneously investigated three ecosystem functions and forage nutrient values following potential species extinction scenarios (dominant species removal, rare species removal, end-member species removal and random species removal) in a Mongolian grassland. ANPP, forage nutrient values, litter decomposition, and soil respiration were measured one and/or two years after plant removal. DNA samples of microorganisms extracted from the soil were subjected to metagenomics analysis. Finally, we calculated the multifunctionality, and examined the relationship of multifunctionality with plant and microorganism diversity. Among ecosystem functions, ANPP and litter decomposition rate decreased under random and rare species extinction scenarios, respectively, and forage quality increased when only dominant species had been removed. Diversity and species composition of soil microorganism were not affected by plant species richness or removal scenario. Only genus-level diversity of bacteria and ANPP were significantly and positively correlated with microbial diversity. Taken together, decreasing species richness of plants and soil organisms rarely impaired multifunctionality. Ecosystem functions were relatively robust to realistic disturbances and species extinction in natural grasslands. However, as each function responded differently to the different sets of species removed, the consequences of a realistic non-random extinction scenario for multiple ecosystem functions should be critical to the management of biodiversity loss caused by different disturbances.     

Seven years of carbon dioxide enrichment, nitrogen fertilization and plant diversity influence arbuscular mycorrhizal fungi in a grassland ecosystem

? We tested the prediction that the abundance and diversity of arbuscular mycorrhizal (AM) fungi are influenced by resource availability and plant community composition by examining the joint effects of carbon dioxide (CO(2) ) enrichment, nitrogen (N) fertilization and plant diversity on AM fungi. ? We quantified AM fungal spores and extramatrical hyphae in 176 plots after 7 yr of treatment with all combinations of ambient or elevated CO(2) (368 or 560 ppm), with or without N fertilization (0 or 4 g Nm(-2) ), and one (monoculture) or 16 host plant species (polyculture) in the BioCON field experiment at Cedar Creek Ecosystem Science Reserve, Minnesota, USA. ? Extramatrical hyphal lengths were increased by CO(2) enrichment, whereas AM spore abundance decreased with N fertilization. Spore abundance, morphotype richness and extramatrical hyphal lengths were all greater in monoculture plots. A structural equation model showed AM fungal biovolume was most influenced by CO(2) enrichment, plant community composition and plant richness, whereas spore richness was most influenced by fungal biovolume, plant community composition and plant richness. ? Arbuscular mycorrhizal fungi responded to differences in host community and resource availability, suggesting that mycorrhizal functions, such as carbon sequestration and soil stability, will be affected by global change.     

内蒙古草原不同基因组大小植物对氮水添加的响应

被子植物基因组大小的种间差异巨大,约为2400倍.基因组大小与植物从细胞核到个体水平的一系列性状密切相关,进而影响植物对环境变化的响应.作为水分和养分共同限制的生态系统,内蒙古草原植物群落对氮素、水分有效性变化的响应具有明显的种间差异,这种差异可能与种间基因组大小不同有关.本研究利用流式细胞术测定了内蒙古典型草原水分、氮素添加实验平台植物的基因组大小,研究了不同基因组大小植物地上净初级生产力(ANPP)和物种丰富度对水分、氮素添加及其交互作用的响应.结果表明:基因组大小显著影响了不同植物ANPP对水分的响应,小基因组植物ANPP对氮水添加响应更敏感,加水和氮水共同添加显著增加了小基因组植物ANPP,而大基因组植物ANPP对所有处理响应均不显著.加氮对大小基因组植物ANPP都无显著影响.大小基因组植物的物种丰富度对氮水添加的响应也均不显著.基因组大小影响内蒙古草原不同植物ANPP对水分增加的响应.作为植物细胞核水平上十分稳定且种间差异巨大的物种性状,将基因组大小引入生态学研究将对全球变化背景下生态系统结构与功能变化研究起到重要作用.     

Nitrogen addition does not reduce the role of spatial asynchrony in stabilising grassland communities

While nitrogen (N) amendment is known to affect the stability of ecological communities, whether this effect is scale‐dependent remains an open question. By conducting a field experiment in a temperate grassland, we found that both plant richness and temporal stability of community biomass increased with spatial scale, but N enrichment reduced richness and stability at the two scales considered. Reduced local‐scale stability under N enrichment arose from N‐induced reduction in population stability, which was partly attributable to the decline in local species richness, as well as reduction in asynchronous local population dynamics across species. Importantly, N enrichment did not alter spatial asynchrony among local communities, which provided similar spatial insurance effects at the larger scale, regardless of N enrichment levels. These results suggest that spatial variability among local communities, in addition to local diversity, may help stabilise ecosystems at larger spatial scales even in the face of anthropogenic environmental changes.     

High land‐use intensity exacerbates shifts in grassland vegetation composition after severe experimental drought

Climate change projections anticipate increased frequency and intensity of drought stress, but grassland responses to severe droughts and their potential to recover are poorly understood. In many grasslands, high land‐use intensity has enhanced productivity and promoted resource‐acquisitive species at the expense of resource‐conservative ones. Such changes in plant functional composition could affect the resistance to drought and the recovery after drought of grassland ecosystems with consequences for feed productivity resilience and environmental stewardship. In a 12‐site precipitation exclusion experiment in upland grassland ecosystems across Switzerland, we imposed severe edaphic drought in plots under rainout shelters and compared them with plots under ambient conditions. We used soil water potentials to scale drought stress across sites. Impacts of precipitation exclusion and drought legacy effects were examined along a gradient of land‐use intensity to determine how grasslands resisted to, and recovered after drought. In the year of precipitation exclusion, aboveground net primary productivity (ANPP) in plots under rainout shelters was ?15% to ?56% lower than in control plots. Drought effects on ANPP increased with drought severity, specified as duration of topsoil water potential ψ < ?100 kPa, irrespective of land‐use intensity. In the year after drought, ANPP had completely recovered, but total species diversity had declined by ?10%. Perennial species showed elevated mortality, but species richness of annuals showed a small increase due to enhanced recruitment. In general, the more resource‐acquisitive grasses increased at the expense of the deeper‐rooted forbs after drought, suggesting that community reorganization was driven by competition rather than plant mortality. The negative effects of precipitation exclusion on forbs increased with land‐use intensity. Our study suggests a synergistic impact of land‐use intensification and climate change on grassland vegetation composition, and implies that biomass recovery after drought may occur at the expense of biodiversity maintenance.     

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.     

Environmental factors covary with plant diversity–productivity relationships among Chinese grassland sites

Aim Our objective was to document the general relationship between plant species richness (SR) and above‐ground net primary productivity (ANPP) at different spatial scales and the environmental influence on this relationship. Location Temperate and alpine grasslands of China. Methods We investigated SR and ANPP at 321 field sites (1355 plots) across the widely distributed temperate and alpine grasslands of China. Ordinary least squares (OLS) regressions were used to test SR–ANPP relationships among site means. Plot‐level data of SR and ANPP were analysed with general linear models (GLMs) and the correlation between SR and ANPP was decomposed into covariance components to test the influence of climatic variables, region, vegetation type and remaining variation among sites on SR, ANPP and their relationship. Results We found positive linear relationships between SR and ANPP among sites in both the alpine and temperate grassland regions and in different grassland vegetation types of these biomes. Environmental gradients such as growing‐season precipitation affected both SR and ANPP in parallel. However, after removing the among‐site environmental variation, residual SR and ANPP were no longer correlated at the pooled within‐site level. Main conclusions The positive SR–ANPP relationship across large‐scale environmental gradients among sites was most likely the result of climatic variables influencing SR and ANPP in parallel. Our results suggest that in China's natural grasslands there is no direct relationship between SR and ANPP, presumably because the pool of available species for local community assembly is large, in contrast to experiments where species pools are artificially reduced.     

Importance of Seasonality for the Response of a Mesic Temperate Grassland to Increased Precipitation Variability and Warming

Timing of precipitation events within the growing season and the non-uniformity of warming might be decisive for alterations in productivity and community composition, with consequences for ecosystem functioning. The responses of aboveground production, community composition, functional group and species evenness to altered intra-annual precipitation variability and their interactions with winter or summer warming were examined in European, mesic temperate grassland. Increased precipitation variability with an induced spring drought resulted in a 17% reduction in ANPP, and late drought reduced ANPP by 18% compared to regular rainfall patterns throughout the entire growing season. Winter warming increased ANPP by 12%, whereas summer warming showed no significant effect on biomass but decreased species richness. The effects of increased precipitation variability and warming on ANPP were independent of each other. Forbs benefited from high precipitation variability with spring drought events, likely due to reduced competitive pressure by decreasing, water stressed grasses. Increased precipitation variability coinciding with higher summer temperatures led to reduced species evenness and likely promoted the establishment of specialists and drought-tolerant species. Seasonality of climatic factors, here early versus late drought events in the high precipitation variability treatments, was important in driving shifts in community composition but not for decreases in ANPP. Non-uniform warming, here winter versus summer, affected the direction of response of both community composition and ANPP. Variability of resources is affecting ecosystem processes and species interactions. Recognition of seasonality and non-uniformity of climatic factors will improve predictions of plant performance and biotic interactions in response to climate change.     

Feedback from plant species change amplifies CO2 enhancement of grassland productivity

Dynamic global vegetation models simulate feedbacks of vegetation change on ecosystem processes, but direct, experimental evidence for feedbacks that result from atmospheric CO₂ enrichment is rare. We hypothesized that feedbacks from species change would amplify the initial CO₂ stimulation of aboveground net primary productivity (ANPP) of tallgrass prairie communities. Communities of perennial forb and C₄ grass species were grown for 5 years along a field CO₂ gradient (250–500 μL L^?1) in central Texas USA on each of three soil types, including upland and lowland clay soils and a sandy soil. CO₂ enrichment increased community ANPP by 0–117% among years and soils and increased the contribution of the tallgrass species Sorghastrum nutans (Indian grass) to community ANPP on each of the three soil types. CO₂‐induced changes in ANPP and Sorghastrum abundance were linked. The slope of ANPP‐CO₂ regressions increased between initial and final years on the two clay soils because of a positive feedback from the increase in Sorghastrum fraction. This feedback accounted for 30–60% of the CO₂‐mediated increase in ANPP on the upland and lowland clay soils during the final 3 years and 1 year of the experiment, respectively. By contrast, species change had little influence on the ANPP‐CO₂ response on the sandy soil, possibly because Sorghastrum increased largely at the expense of a functionally similar C₄ grass species. By favoring a mesic C₄ tall grass, CO₂ enrichment approximately doubled the initial enhancement of community ANPP on two clay soils. The CO₂‐stimulation of grassland productivity may be significantly underestimated if feedbacks from plant community change are not considered.