Defoliation and competition effects in a productivity gradient for a semiarid Mediterranean annual grassland community

Grazing and competition are two main factors shaping range plant communities; however, few studies have investigated their interaction. The current study aimed to investigate the effects of defoliation, competition and their interaction on production of annual grasses in semiarid Mediterranean areas. Competition treatments (absence/presence of neighbors) were combined with three defoliation intensities (0%, 30% and 60%) in a complete factorial design. Competition significantly reduced grass biomass. However, the role of competition was eliminated under heavy defoliation or under dry growth conditions. Defoliation showed variable results on final biomass (FB) and cumulative biomass (CB). While heavy defoliation (60% clipping intensity) reduced grass FB down to 80% during the two growing seasons, light defoliation (30%) significantly increased CB. Results showed that competition may limit the direct effect of defoliation on dominant grass species. Further, the relationship between site productivity and competition effect was best explained by a negative linear model. This hypothesized model may suggest that facilitation and competition alternatively affect grassland communities along a productivity gradient. The results suggest that light grazing may sustain or even enhance grassland productivity. The results also indicated the suitability of annual grass species to re-vegetate degraded rangeland in semi-arid climate. Further, optimum grazing practices to conserve biodiversity of Avena grassland may involve moderate stocking rate.     

Fungal Symbionts Alter Plant Drought Response

Grassland productivity is often primarily limited by water availability, and therefore, grasslands may be especially sensitive to climate change. Fungal symbionts can mediate plant drought response by enhancing drought tolerance and avoidance, but these effects have not been quantified across grass species. We performed a factorial meta-analysis of previously published studies to determine how arbuscular mycorrhizal (AM) fungi and endophytic fungal symbionts affect growth of grasses under drought. We then examined how the effect of fungal symbionts on plant growth was influenced by biotic (plant photosynthetic pathway) and abiotic (level of drought) factors. We also measured the phylogenetic signal of fungal symbionts on grass growth under control and drought conditions. Under drought conditions, grasses colonized by AM fungi grew larger than those without mycorrhizal symbionts. The increased growth of grasses conferred from fungal symbionts was greatest at the lowest soil moisture levels. Furthermore, under both drought and control conditions, C3 grasses colonized by AM fungi grew larger than C3 grasses without symbionts, but the biomass of C4 grasses was not affected by AM fungi. Endophytes did not increase plant biomass overall under any treatment. However, there was a phylogenetically conserved increase in plant biomass in grasses colonized by endophytes. Grasses and their fungal symbionts seem to interact within a context-dependent symbiosis, varying with biotic and abiotic conditions. Because plant–fungal symbioses significantly alter plant drought response, including these responses could improve our ability to predict grassland functioning under global change.     

The type of competition modulates the ecophysiological response of grassland species to elevated CO2 and drought

The effects of elevated CO₂ and drought on ecophysiological parameters in grassland species have been examined, but few studies have investigated the effect of competition on those parameters under climate change conditions. The objective of this study was to determine the effect of elevated CO₂ and drought on the response of plant water relations, gas exchange, chlorophyll a fluorescence and aboveground biomass in four grassland species, as well as to assess whether the type of competition modulates that response. Elevated CO₂ in well‐watered conditions increased aboveground biomass by augmenting CO₂ assimilation. Drought reduced biomass by reducing CO₂ assimilation rate via stomatal limitation and, when drought was more severe, also non‐stomatal limitation. When plants were grown under the combined conditions of elevated CO₂ and drought, drought limitation observed under ambient CO₂ was reduced, permitting higher CO₂ assimilation and consequently reducing the observed decrease in aboveground biomass. The response to climate change was species‐specific and dependent on the type of competition. Thus, the response to elevated CO₂ in well‐watered grasses was higher in monoculture than in mixture, while it was higher in mixture compared to monoculture for forbs. On the other hand, forbs were more affected than grasses by drought in monoculture, while in mixture the negative effect of drought was higher in grasses than in forbs, due to a lower capacity to acquire water and mineral nutrients. These differences in species‐level growth responses to CO₂ and drought may lead to changes in the composition and biodiversity of the grassland plant community in future climate conditions.     

Water stress in grasslands: dynamic responses of plants and insect herbivores

Global climate change is altering precipitation patterns. The effect of water stress on plant–herbivore interactions is poorly understood even though this is a primary ecological interaction that will be altered by climate change. This is especially true for grasslands where water is often limiting. In this study we manipulated water inputs in open grassland plots (1 m²) during a severe drought and assessed plant and insect herbivore responses. There were two watering treatments: ambient and supplemented. Supplemented plots received water weekly in amounts that mimicked average seasonal rainfall. For plants, we were interested in how water input affected protein and digestible carbohydrate content; previous studies predicted water stress would increase the concentration of these two nutrients. Grasshoppers are the dominant insect herbivores in grasslands and we assessed their responses to water inputs by measuring abundance and diversity. Previous studies suggested grasshoppers would prefer water‐stressed plots. Protein and carbohydrate content in bulk grass and forb samples, plus plant biomass and diversity, were measured monthly (May–August). Immediately prior to harvesting plant tissue, we counted and identified individual grasshoppers in each plot. Grass biomass was reduced with water stress, but macronutrient content and species diversity were unaffected. After three months water‐stressed forbs were less protein biased, and diverse, relative to watered forbs; forb biomass was indistinguishable between treatments. Grasshopper abundance and diversity were lower in water‐stressed plots as the season progressed. However, grasshopper‐feeding biology mattered: densities of mixed‐feeders and grass‐feeders, but not forb‐specialists, decreased over time in water‐stressed plots, but not in water supplemented plots. Our results demonstrate the importance of focusing on plant and insect herbivore functional groups and provide valuable new data that can be incorporated into models to explore the effects of global climate change in greater detail.     

Selective grazing modifies previously anticipated responses of plant community composition to elevated CO2 in a temperate grassland

Our limited understanding of terrestrial ecosystem responses to elevated CO₂ is a major constraint on predicting the impacts of climate change. A change in botanical composition has been identified as a key factor in the CO₂ response with profound implications for ecosystem services such as plant production and soil carbon storage. In temperate grasslands, there is a strong consensus that elevated CO₂ will result in a greater physiological stimulus to growth in legumes and to a lesser extent forbs, compared with C3 grasses, and the presumption this will lead in turn to a greater proportion of these functional groups in the plant community. However, this view is based on data mainly collected in experiments of three or less years in duration and not in experiments where defoliation has been by grazing animals. Grazing is, however, the most common management of grasslands and known in itself to influence botanical composition. In a long‐term Free Air Carbon Dioxide Enrichment (FACE) experiment in a temperate grassland managed with grazing animals (sheep), we found the response to elevated CO₂ in plant community composition in the first 5 years was consistent with the expectation of increased proportions of legumes and forbs. However, in the longer term, these differences diminished so that the proportions of grasses, legumes and forbs were the same under both ambient and elevated CO₂. Analysis of vegetation before and after each grazing event showed there was a sustained disproportionately greater removal (‘apparent selection’) of legumes and forbs by the grazing animals. This bias in removal was greater under elevated CO₂ than ambient CO₂. This is consistent with sustained faster growth rates of legumes and forbs under elevated CO₂ being countered by selective defoliation, and so leading to little difference in community composition.     

Biomass production potential of grasslands in the oak savanna region of Minnesota, USA

Biomass harvested from grasslands formerly used for forage production or set aside for conservation has been identified as a potential source of bioenergy feedstocks. Our objective was to characterize yield and chemical composition of biomass harvested from existing grasslands in the oak savanna region of Minnesota and to determine whether aggregated soil properties and grassland type influence biomass yield and feedstock properties. The influence of soil type and dominant functional plant groups on biomass yield, theoretical ethanol yield, and mineral, ash, and lignocellulosic concentration was measured on biomass harvested from 32 grassland sites. Soils with high productivity ratings, as measured by the Minnesota Crop Productivity Index, produced 36 % more biomass than lower quality soils. Grasslands dominated by warm-season species produced 18 % more biomass than those dominated by cool-season species, when measured after senescence during the late-fall harvest time. Biomass harvested from sites dominated by cool-season grasses had higher N, Mg and Cl concentrations than those dominated by warm-season grasses, suggesting that such grasslands could have lower efficiency in thermochemical conversion processes and that repeated harvesting from such grasslands could remove nutrients from the systems. In addition, glucose and xylose concentrations were slightly higher in biomass from sites dominated by warm-season grasses, which resulted in an estimated additional 12 L?Mg^?1 of ethanol over those dominated by cool-season grasses. Combined with the greater yields, warm-season grasslands could produce an additional 376 L?ha^?1 year^?1.     

Recovery of plant species composition and ecosystem function after cessation of grazing in a Mediterranean grassland

Short- and long-term changes in species composition, plant biomass production, and litter decomposition after cessation of grazing were examined in a Mediterranean grassland with high dominance of annual species and strong seasonality in biomass production. Short-term changes were assessed during three consecutive years in plots previously exposed to different grazing pressures and compared to plots in long-term (30–40 years) exclosures. Short-term cessation of grazing led in the short-term to an increase in relative biomass of annual crucifers and tall annual and perennial grasses, while biomass of annual legumes, annual thistles and short annual grasses decreased. Consequently, similarity increased between vegetation recently excluded from grazing and vegetation in long-term protected plots. Our research showed that in systems with high dominance of grasses and annual species, the rapid changes in plant species composition that occur after grazing cessation were associated with a fast recovery of the potential for biomass production to levels found in long-term protected plots, while litter decomposition rate did not change even after long-term cessation of grazing. Moreover, previous history of grazing did not affect plant litter decomposition, despite higher litter quality in grazed treatments. This study provides new insights about the processes involved in the diverse responses of ecosystem functions resulting from shifts in species composition associated with grazing cessation and land use change in Mediterranean grasslands.     

Spatial patterns of soil and ecosystem respiration regulated by biological and environmental variables along a precipitation gradient in semi-arid grasslands in China

Precipitation is a key environmental factor in determining ecosystem structure and function. Knowledge of how soil and ecosystem respiration responds to climate change (e.g., precipitation) and human activities (e.g., grazing or clipping) is crucial for assessing the impacts of climate change on terrestrial ecosystems and for improving model simulations and predictions of future global carbon (C) cycling in response to human activities. In this study, we examined the spatial patterns of soil and ecosystem respiration along a precipitation gradient from 167.7 to 398.1 mm in a semi-arid grassland. Our results showed that soil and ecosystem respiration increased linearly with increasing mean annual precipitation. The trends were similar to those of shoot biomass, litter and soil total C content along the precipitation gradient. Our results indicated that precipitation was the primary controlling factor in determining the spatial pattern of soil and ecosystem respiration in semi-arid grasslands in China. The linear/nonlinear relationships in this study describing the variations of the ecosystem carbon process with precipitation can be useful for model development, parameterization and validation at the regional scale to improve predictions of how carbon processes in grasslands respond to climate change, land use and grassland management.     

Lasting effects of climate disturbance on perennial grassland above‐ground biomass production under two cutting frequencies

Climate extremes can ultimately reshape grassland services such as forage production and change plant functional type composition. This 3‐year field research studied resistance to dehydration and recovery after rehydration of plant community and plant functional types in an upland perennial grassland subjected to climate and cutting frequency (Cut+, Cut?) disturbances by measuring green tissue percentage and above‐ground biomass production (ANPP). In year 1, a climate disturbance gradient was applied by co‐manipulating temperature and precipitation. Four treatments were considered: control and warming‐drought climatic treatment, with or without extreme summer event. In year 2, control and warming‐drought treatments were maintained without extreme. In year 3, all treatments received ambient climatic conditions. We found that the grassland community was very sensitive to dehydration during the summer extreme: aerial senescence reached 80% when cumulated climatic water balance fell to ?156 mm and biomass declined by 78% at the end of summer. In autumn, canopy greenness and biomass totally recovered in control but not in the warming‐drought treatment. However ANPP decreased under both climatic treatments, but the effect was stronger on Cut+ (?24%) than Cut? (?15%). This decline was not compensated by the presence of three functional types because they were negatively affected by the climatic treatments, suggesting an absence of buffering effect on grassland production. In the following 2 years, lasting effects of climate disturbance on ANPP were observable. The unexpected stressful conditions of year 3 induced a decline in grassland production in the Cut+ control treatment. The fact that this treatment cumulated higher (45%) N export over the 3 years suggests that N plays a key role in ANPP stability. As ANPP in this mesic perennial grassland did not show engineering resilience, long‐term experimental manipulation is needed. Infrequent mowing appears more appropriate for sustaining grassland ANPP under future climate extremes.     

Clipping defoliation and nitrogen addition shift competition between a C3 grass (Leymus chinensis) and a C4 grass (Hemarthria altissima)

• Human‐induced disturbances, including grazing and clipping, that cause defoliation are common in natural grasslands. Plant functional type differences in the ability to compensate for this tissue loss may influence interspecific competition.
• To explore the effects of different intensities of clipping and nitrogen (N) addition on compensatory growth and interspecific competition, we measured accumulated aboveground biomass (AGB), belowground biomass (BGB), tiller number, non‐structural carbohydrates concentrations and leaf gas exchange parameters in two locally co‐occurring species (the C₃ grass Leymus chinensis and the C₄ grass Hemarthria altissima) growing in monoculture and in mixture.
• For both grasses, the clipping treatment had significant impacts on the accumulated AGB, and the 40% clipping treatment had the largest effect. BGB gradually decreased with increasing defoliation intensity. Severe defoliation caused a significant increase in tiller number. Stored carbohydrates in the belowground biomass were mobilised and transported aboveground for the growth of new leaves to compensate for clipping‐induced injury. The net CO₂ assimilation rate (A) of the remaining leaves increased with clipping intensity and peaked under clipping intensities of 20% or 40%. Nitrogen addition, at a rate of 10 g·N·m^?2·year^?1, enhanced A of the remaining leaves and non‐structural carbohydrate concentrations, which benefited plant compensatory growth, especially for the C₃ grass. Under the mixed planting conditions, the clipping and N addition treatments lowered the competitive advantage of the C₄ grass.
• The results suggest that a combination of defoliation and N deposition have the potential to benefit the coexistence of C₃ and C₄ grasses.

     

干旱半干旱草地生态系统与土壤水分关系研究进展

研究干旱半干旱草地生态系统与土壤水分关系和相互作用机理对于揭示草地生态系统稳定性及其水土关键要素的变化过程具有重要意义。从不同界面、不同尺度综述了草地生态系统对土壤水分的影响及草地生态系统的响应与适应机制,总结了草地生态系统与土壤水分关系模型研究的相关进展,并分析了气候变化对草地生态系统和土壤水分关系的影响。草地生态系统通过影响水文过程和生态过程来影响土壤水分,土壤水分在植物生长发育、形态、生理生态过程、种间关系、群落组成和结构以及草地生态系统功能等方面对草地生态系统产生影响;充分揭示草地生态系统-土壤水分相互作用机理是模型研究的关键;气候变化对草地生态系统植物与土壤水分关系具有重要影响。今后应加强以下研究:1)开展草地不同优势种和植物功能型与土壤水分关系的研究,找出能反映植物对土壤水分响应的性状指标,阈值响应点及适应机制;2)注重对不同时间和空间尺度上的转换和比较;3)加强个体、群体和生态系统尺度草地植物生长模型的研究及其与土壤-植被-大气水分传输模型的耦合;4)加强草地生态系统与土壤水分关系对气候变化响应的研究。     

Site‐specific responses of native and exotic species to disturbances in a mesic grassland community

Abstract. Grassland communities are increasingly recognized as disturbance‐dependent ecosystems, yet there are few replicated, multi‐site studies documenting vegetation responses to varying frequencies and types of grassland disturbance. Even so, land managers frequently manipulate disturbance regimes in an attempt to favour native grassland plants over exotic species. We conducted a factorial experiment testing three frequencies of clipping combined with litter accumulation, litter removal, and soil disturbance within the highly threatened California coastal prairie plant community. We monitored the response of native/exotic, grass/forb plant guilds once a year for four years. More frequent clipping reduced cover of exotic grasses and favoured exotic forbs, whereas native species were largely unaffected by clipping frequency. Litter accumulation, litter removal, and soil disturbance did not affect vegetation composition. Effects of litter accumulation may take longer than our experiment allowed, and soil disturbance due to our treatments was not sufficiently strong to show consistent effects relative to mammalian soil disturbance. Treatment response of some plant guilds differed among sites, highlighting the importance of replicating experiments at several sites before recommending conservation management practices.     

Adaptive phenotypic plasticity of Pseudoroegneria spicata: response of stomatal density, leaf area and biomass to changes in water supply and increased temperature

Background and Aims

Changes in rainfall and temperature brought about through climate change may affect plant species distribution and community composition of grasslands. The primary objective of this study was to test how manipulation of water and temperature would influence the plasticity of stomatal density and leaf area of bluebunch wheatgrass, Pseudoroegneria spicata. It was hypothesized that: (1) an increased water supply will increase biomass and leaf area and decrease stomatal density, while a reduced water supply will cause the opposite effect; (2) an increase in temperature will reduce biomass and leaf area and increase stomatal density; and (3) the combinations of water and temperature treatments can be aligned along a stress gradient and that stomatal density will be highest at high stress.

Methods

The three water supply treatments were (1) ambient, (2) increased approx. 30 % more than ambient through weekly watering and (3) decreased approx. 30 % less than ambient by rain shades. The two temperature treatments were (1) ambient and (2) increased approx. 1–3 °C by using open-top chambers. At the end of the second experimental growing season, above-ground biomass was harvested, oven-dried and weighed, tillers from bluebunch wheatgrass plants sampled, and the abaxial stomatal density and leaf area of tillers were measured.

Key Results

The first hypothesis was partially supported – reducing water supply increased stomatal density, but increasing water supply reduced leaf area. The second hypothesis was rejected. Finally, the third hypothesis could not be fully supported – rather than a linear response there appears to be a parabolic stomatal density response to stress.

Conclusions

Overall, the abaxial stomatal density and leaf area of bluebunch wheatgrass were plastic in their response to water and temperature manipulations. Although bluebunch wheatgrass has the potential to adapt to changing climate, the grass is limited in its ability to respond to a combination of reduced water and increased temperature.Key words: Bluebunch wheatgrass, Pseudoroegneria spicata, biomass, climate change, grassland, open top chamber, rain shade, stomata     

Defoliation of Centaurea solstitialis Stimulates Compensatory Growth and Intensifies Negative Effects on Neighbors

Efforts to arrest the spread of invasive weeds with herbivory may be hindered by weak effects of the herbivores or strong compensatory responses of the invaders. We conducted a greenhouse experiment to study the effects of defoliation and soil fungi on competition between the invasive weed Centaurea solstitialis and C. solstitialis and Avena barbata, a naturalized Eurasian annual grass, and Nassella pulchra, a native California bunchgrass. Surprisingly, considering the explosive invasion of grasslands by C. solstitialis, Avena and Nassella were strong competitors and reduced the invader’s biomass by 80.2% and 80.1% over all defoliation and soil fungicide treatments, respectively. However, our experiments were conducted in artificial environments where competition was probably accentuated. When fungicide was applied to the soil, the biomass of C. solstitialis was reduced in all treatment combinations, but reduction in the biomass of the invader had no corollary impact on the grasses. There was no overall effect of defoliation on the final biomass of C. solstitialis as the invader compensated fully for severe clipping. In fact, the directional trend of the clipping effect was +6.4% over all treatments after eight weeks. A significant neighbor × soil fungicide × clipping effect suggested that the compensatory response was the strongest without soil fungicide and when C. solstitialis was alone (+ 19%). Our key finding was that the compensatory response of C. solstitialis in all treatments was associated with an increase in the weed’s negative effects on Nassella and Avena – there was a significant decrease in the total biomass of both grasses and the reproductive biomass of Avena in pots with clipped C. solstitialis. Our results were obtained in controlled conditions that may have been conducive to compensatory growth, but they suggest the existence of mechanisms that may allow C. solstitialis, like other Centaurea species, to resist herbivory.     

Productivity responses to altered rainfall patterns in a C<Subscript>4</Subscript>-dominated grassland

Rainfall variability is a key driver of ecosystem structure and function in grasslands worldwide. Changes in rainfall patterns predicted by global climate models for the central United States are expected to cause lower and increasingly variable soil water availability, which may impact net primary production and plant species composition in native Great Plains grasslands. We experimentally altered the timing and quantity of growing season rainfall inputs by lengthening inter-rainfall dry intervals by 50%, reducing rainfall quantities by 30%, or both, compared to the ambient rainfall regime in a native tallgrass prairie ecosystem in northeastern Kansas. Over three growing seasons, increased rainfall variability caused by altered rainfall timing with no change in total rainfall quantity led to lower and more variable soil water content (0–30 cm depth), a ~10% reduction in aboveground net primary productivity (ANPP), increased root to shoot ratios, and greater canopy photon flux density at 30 cm above the soil surface. Lower total ANPP primarily resulted from reduced growth, biomass and flowering of subdominant warm-season C₄ grasses while productivity of the dominant C₄ grass Andropogon gerardii was relatively unresponsive. In general, vegetation responses to increased soil water content variability were at least equal to those caused by imposing a 30% reduction in rainfall quantity without altering the timing of rainfall inputs. Reduced ANPP most likely resulted from direct effects of soil moisture deficits on root activity, plant water status, and photosynthesis. Altered rainfall regimes are likely to be an important element of climate change scenarios in this grassland, and the nature of interactions with other climate change elements remains a significant challenge for predicting ecosystem responses to climate change.     

Plant and arthropod community sensitivity to rainfall manipulation but not nitrogen enrichment in a successional grassland ecosystem

Grasslands provide many ecosystem services including carbon storage, biodiversity preservation and livestock forage production. These ecosystem services will change in the future in response to multiple global environmental changes, including climate change and increased nitrogen inputs. We conducted an experimental study over 3 years in a mesotrophic grassland ecosystem in southern England. We aimed to expose plots to rainfall manipulation that simulated IPCC 4th Assessment projections for 2100 (+15 % winter rainfall and ?30 % summer rainfall) or ambient climate, achieving +15 % winter rainfall and ?39 % summer rainfall in rainfall-manipulated plots. Nitrogen (40 kg ha^?1 year^?1) was also added to half of the experimental plots in factorial combination. Plant species composition and above ground biomass were not affected by rainfall in the first 2 years and the plant community did not respond to nitrogen enrichment throughout the experiment. In the third year, above-ground plant biomass declined in rainfall-manipulated plots, driven by a decline in the abundances of grass species characteristic of moist soils. Declining plant biomass was also associated with changes to arthropod communities, with lower abundances of plant-feeding Auchenorrhyncha and carnivorous Araneae indicating multi-trophic responses to rainfall manipulation. Plant and arthropod community composition and plant biomass responses to rainfall manipulation were not modified by nitrogen enrichment, which was not expected, but may have resulted from prior nitrogen saturation and/or phosphorus limitation. Overall, our study demonstrates that climate change may in future influence plant productivity and induce multi-trophic responses in grasslands.     

Ecological conditions that determine when grazing stimulates grass production

Summary We report the results of a pot experiment that examined the effects of three ecologically important factors controlling plant growth rates in savanna grasslands: defoliation, soil nitrogen and soil water availability. The experiment was conducted in the Amboseli region in east Africa, and was designed to simulate natural conditions as far as possible, using local soils and a grass species that is heavily grazed by abundant large herbivores. Productivity by different plant components was reduced, stimulated or unchanged by defoliation, depending on specific watering and fertilization treatments. Total above-ground production was stimulated by defoliation and was maximized at moderate clipping intensities, but this was statistically significant only when plants were watered infrequently (every 8 days), and most important, periods between clipping events were extended (at least 24 days). Under these conditions, plant growth rates were limited by water availability at the time of clipping, and soil water conserved in clipped, compared to unclipped plants. Within a given fertilization treatment, whole-plant production was never stimulated by defoliation because root growth was unaffected or inhibited by clipping. However, when fertilization was coupled to defoliation, as they are in the field, whole-plant production by fertilized and moderately clipped plants exceeded production by infertilized, unclipped plants. Under this interpretation, maximum whole-plant production coincided with optimum conditions for herbivores (maximum nitrogen concentration in grass leaves) when watering was frequent, and plants were moderately defoliated. However, these conditions were not the same as those that maximized relative above-ground stimulation of growth (infrequent watering and clipping).The results indicate that above-ground grass production can be stimulated by grazing, and when that is likely to occur. However, the results emphasize that plant production responses to defoliation can vary widely, contigent upon a complex interaction of ecological factors.     

Plant community responses to increased precipitation and belowground litter addition: Evidence from a 5‐year semiarid grassland experiment

Global climate change is predicted to stimulate primary production and consequently increases litter inputs. Changing precipitation regimes together with enhanced litter inputs may affect plant community composition and structure, with consequent influence on diversity and ecosystem functioning. Responses of plant community to increased precipitation and belowground litter addition were examined lasting 5 years in a semiarid temperate grassland of northeastern China. Increased precipitation enhanced community species richness and abundance of annuals by 16.8% and 44%, but litter addition suppressed them by 25% and 54.5% after 5 years, respectively. During the study period, perennial rhizome grasses and forbs had consistent negative relationship under ambient plots, whereas positive relationship between the two functional groups was found under litter addition plots after 5 years. In addition, increased precipitation and litter addition showed significant interaction on community composition, because litter addition significantly increased biomass and abundance of rhizome grasses under increased precipitation plots but had no effect under ambient precipitation levels. Our findings emphasize the importance of water availability in modulating the responses of plants community to potentially enhanced litter inputs in the semiarid temperate grassland.     

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.     

From facilitation to competition: temperature‐driven shift in dominant plant interactions affects population dynamics in seminatural grasslands

Biotic interactions are often ignored in assessments of climate change impacts. However, climate‐related changes in species interactions, often mediated through increased dominance of certain species or functional groups, may have important implications for how species respond to climate warming and altered precipitation patterns. We examined how a dominant plant functional group affected the population dynamics of four co‐occurring forb species by experimentally removing graminoids in seminatural grasslands. Specifically, we explored how the interaction between dominants and subordinates varied with climate by replicating the removal experiment across a climate grid consisting of 12 field sites spanning broad‐scale temperature and precipitation gradients in southern Norway. Biotic interactions affected population growth rates of all study species, and the net outcome of interactions between dominants and subordinates switched from facilitation to competition with increasing temperature along the temperature gradient. The impacts of competitive interactions on subordinates in the warmer sites could primarily be attributed to reduced plant survival. Whereas the response to dominant removal varied with temperature, there was no overall effect of precipitation on the balance between competition and facilitation. Our findings suggest that global warming may increase the relative importance of competitive interactions in seminatural grasslands across a wide range of precipitation levels, thereby favouring highly competitive dominant species over subordinate species. As a result, seminatural grasslands may become increasingly dependent on disturbance (i.e. traditional management such as grazing and mowing) to maintain viable populations of subordinate species and thereby biodiversity under future climates. Our study highlights the importance of population‐level studies replicated under different climatic conditions for understanding the underlying mechanisms of climate change impacts on plants.