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.     

Elevated CO2 and water addition enhance nitrogen turnover in grassland plants with implications for temporal stability

Temporal variation in soil nitrogen (N) availability affects growth of grassland communities that differ in their use and reuse of N. In a 7‐year‐long climate change experiment in a semi‐arid grassland, the temporal stability of plant biomass production varied with plant N turnover (reliance on externally acquired N relative to internally recycled N). Species with high N turnover were less stable in time compared to species with low N turnover. In contrast, N turnover at the community level was positively associated with asynchrony in biomass production, which in turn increased community temporal stability. Elevated CO₂ and summer irrigation, but not warming, enhanced community N turnover and stability, possibly because treatments promoted greater abundance of species with high N turnover. Our study highlights the importance of plant N turnover for determining the temporal stability of individual species and plant communities affected by climate change.     

Potential trajectories of old‐growth Neotropical forest functional composition under climate change

Quantifying relationships between plant functional traits and abiotic gradients is valuable for evaluating potential responses of forest communities to climate change. However, the trajectories of change expected to occur in tropical forest functional characteristics as a function of future climate variation are largely unknown. We modeled community level trait values of Costa Rican rain forests as a function of current and future climate, and quantified potential changes in functional composition. We calculated per‐plot community weighted mean (CWM) trait values for leaf area (LA), specific leaf area (SLA), leaf dry matter content (LDMC), leaf nitrogen (N) and phosphorus (P) content, and wood basic specific gravity (WSG), for tree and palm species in 127 0.25 ha plots. We modeled the response of CWM traits to current temperature and precipitation gradients using generalized additive modeling. We then predicted and mapped CWM traits values under current and future climate, and quantified potential changes under a global warming scenario (RCP8.5, year 2050). We calculated the area within the multi trait functional space occupied by forest plots under both current and future climate, and determined potential changes in functional space occupied by forest plots. Overall, precipitation predicted CWM traits better than temperature. Models indicated increases in CWM SLA, N and P, and a decrease in CWM LDMC under climate change. Lowland forest communities converged on a single direction of change towards more acquisitive CWM trait values, indicating a change in forest functional composition resulting from a changed climate. Functional space occupied by forest plots was reduced by 50% under the future climate. Functional composition changes may have further effects on forests ecosystem services. Assessing functional trait spatial‐gradients can help bridge the gap between species‐based biogeography and biogeochemical approaches to strengthen biodiversity and ecosystem services conservation efforts.     

Stability in ecosystem functioning across a climatic threshold and contrasting forest regimes

Classical ecological theory predicts that changes in the availability of essential resources such as nitrogen should lead to changes in plant community composition due to differences in species-specific nutrient requirements. What remains unknown, however, is the extent to which climate change will alter the relationship between plant communities and the nitrogen cycle. During intervals of climate change, do changes in nitrogen cycling lead to vegetation change or do changes in community composition alter the nitrogen dynamics? We used long-term ecological data to determine the role of nitrogen availability in changes of forest species composition under a rapidly changing climate during the early Holocene (16k to 8k cal. yrs. BP). A statistical computational analysis of ecological data spanning 8,000 years showed that secondary succession from a coniferous to deciduous forest occurred independently of changes in the nitrogen cycle. As oak replaced pine under a warming climate, nitrogen cycling rates increased. Interestingly, the mechanism by which the species interacted with nitrogen remained stable across this threshold change in climate and in the dominant tree species. This suggests that changes in tree population density over successional time scales are not driven by nitrogen availability. Thus, current models of forest succession that incorporate the effects of available nitrogen may be over-estimating tree population responses to changes in this resource, which may result in biased predictions of future forest dynamics under climate warming.     

Soil heterogeneity buffers community response to climate change in species‐rich grassland

Climate change impacts on vegetation are mediated by soil processes that regulate rhizosphere water balance, nutrient dynamics, and ground‐level temperatures. For ecosystems characterized by high fine‐scale substrate heterogeneity such as grasslands on poorly developed soils, effects of climate change on plant communities may depend on substrate properties that vary at the scale of individuals (<m²), leading to fine‐scale shifts in community structure that may go undetected at larger scales. Here, we show in a long‐running climate experiment in species‐rich limestone grassland in Buxton, England (UK), that the resistance of the community to 15‐year manipulations of temperature and rainfall at the plot scale (9 m²) belies considerable community reorganization at the microsite (100 cm²) scale. In individual models of the abundance of the 25 most common species with respect to climate treatment and microsite soil depth, 13 species exhibited significant soil depth affinities, and nine of these have shifted their position along the depth gradient in response to one or more climate treatments. Estimates of species turnover across the depth gradient reviewed in relation to measurements of water potential, nitrogen supply, pH, and community biomass suggest that communities of shallow microsites are responding directly to microenvironmental changes induced by climate manipulation, while those of the deepest microsites are shifting in response to changes in competitive interference from more nutrient‐demanding species. Moreover, for several species in summer drought and winter heated treatments, climate response in deep microsites was opposite that of shallow microsites, suggesting microsite variation is contributing to community stability at the whole‐plot level. Our study thus demonstrates a strong link between community dynamics and substrate properties, and suggests ecosystems typified by fine‐scale substrate heterogeneity may possess a natural buffering capacity in the face of climate change.     

Impacts of nitrogen deposition on terrestrial plant diversity: a meta-analysis in China

Aims With the global atmospheric nitrogen (N) deposition increasing, the effect of N deposition on terrestrial plant diversity has been widely studied. Some studies have reviewed the effects of N deposition on plant species diversity; however, all studies addressed the effects of N deposition on plant community focused on species richness in specific ecosystem. There is a need for a systematic meta-analysis covering multiple dimensions of plant diversity in multiple climate zones and ecosystems types. Our goal was to quantify changes in species richness, evenness and uncertainty in plant communities in response to N addition across different environmental and experimental contexts.     

模拟氮沉降和降雨变化对贝加尔针茅草原土壤细菌群落结构的影响

研究氮沉降和降雨变化对土壤细菌群落结构的影响,对未来预测多个气候变化因子对草地生态系统影响的交互作用具有重要意义。以施氮和灌溉分别模拟氮沉降和降雨增加,采用高通量测序技术,研究8个氮添加水平(0、15、30、50、100、150、200、300kg N hm-2a-1)和2个水分添加水平(不灌溉、模拟夏季增雨100 mm灌溉)对土壤细菌群落结构的影响。结果表明,氮素和水分输入增加后,土壤细菌群落组成、丰度均显著变化(P0.05)。在群落中占主导的细菌门类有疣微菌门Verrucomicrobia(30.61%—48.51%)、变形菌门Proteobacteria(21.37%—29.97%)、酸杆菌门Acidobacteria(9.54%—20.67%)和拟杆菌门Bacteroidetes(4.96%—9.74%)。在常规降雨和水分添加两种条件下,随着氮添加水平的增加,占主导的细菌门类(相对丰度1%)表现出不同的变化趋势。疣微菌门相对丰度在常规降雨N100—N300条件下显著降低,但在氮素和水分同时添加条件下随氮添加水平升高而逐渐升高,在N200—N300时显著升高。变形菌门和拟杆菌门相对丰度在常规降雨高氮添加条件下呈升高趋势,但在水分添加时却无明显变化。酸杆菌门相对丰度在常规降雨高氮添加条件下升高,但在水分添加后呈明显下降趋势。放线菌门Actinobacteria相对丰度在常规降雨N100—N300条件下显著升高,但在水分添加后高氮添加时显著降低。厚壁菌门Firmicutes相对丰度在常规降雨条件下无显著变化,但在水分和高氮添加条件下降低。浮霉菌门Planctomycetes相对丰度在两种不同的水分添加条件下均呈先升高后降低的趋势。氮素和水分添加对土壤细菌群落结构的变化存在明显的互作效应(P0.0001)。在不同氮素和水分输入条件下共有19个土壤细菌门类相对丰度有显著差异。土壤细菌群落结构的变化主要来自于疣微菌门和酸杆菌门的相对丰度变化,两者可作为土壤细菌群落结构变化的指示种。综上,氮素和水分添加显著改变了土壤细菌群落结构,氮素和水分对土壤细菌不同门类相对丰度变化存在明显的互作效应。     

Links between soil microbial communities and plant traits in a species‐rich grassland under long‐term climate change

Climate change can influence soil microorganisms directly by altering their growth and activity but also indirectly via effects on the vegetation, which modifies the availability of resources. Direct impacts of climate change on soil microorganisms can occur rapidly, whereas indirect effects mediated by shifts in plant community composition are not immediately apparent and likely to increase over time. We used molecular fingerprinting of bacterial and fungal communities in the soil to investigate the effects of 17 years of temperature and rainfall manipulations in a species‐rich grassland near Buxton, UK. We compared shifts in microbial community structure to changes in plant species composition and key plant traits across 78 microsites within plots subjected to winter heating, rainfall supplementation, or summer drought. We observed marked shifts in soil fungal and bacterial community structure in response to chronic summer drought. Importantly, although dominant microbial taxa were largely unaffected by drought, there were substantial changes in the abundances of subordinate fungal and bacterial taxa. In contrast to short‐term studies that report high resistance of soil fungi to drought, we observed substantial losses of fungal taxa in the summer drought treatments. There was moderate concordance between soil microbial communities and plant species composition within microsites. Vector fitting of community‐weighted mean plant traits to ordinations of soil bacterial and fungal communities showed that shifts in soil microbial community structure were related to plant traits representing the quality of resources available to soil microorganisms: the construction cost of leaf material, foliar carbon‐to‐nitrogen ratios, and leaf dry matter content. Thus, our study provides evidence that climate change could affect soil microbial communities indirectly via changes in plant inputs and highlights the importance of considering long‐term climate change effects, especially in nutrient‐poor systems with slow‐growing vegetation.     

贝加尔针茅草原土壤真菌群落结构对氮素和水分添加的响应

研究氮沉降和降雨变化对土壤真菌群落结构的互作效应,对未来预测多个气候变化因子对草地生态系统的交互作用具有重要意义。以施氮和灌溉模拟氮沉降和降雨增加,采用裂区设计,应用高通量测序技术,研究8个氮添加水平(0、15、30、50、100、150、200、300 kg N hm~(-2)a~(-1))和2个水分添加水平(不灌溉、模拟夏季增雨100 mm灌溉)对土壤真菌群落结构的影响。结果表明,氮素和水分添加后,土壤真菌群落中占优势的门类分别为接合菌门Zygomycota(22.0%—48.9%)、担子菌门Basidiomycota(7.8%—18.5%)、子囊菌门Ascomycota(9.4%—20.1%)、球囊菌门Glomeromycota(0.7%—3.1%)、壶菌门Chytridiomycota(0.1%—1.3%)。常规降雨条件下,随着氮添加水平升高,接合菌门相对丰度呈现出先升高后降低的趋势,N50处理最高;子囊菌门相对丰度在高氮添加时(N100—N300)呈升高趋势。而在氮素和水分同时添加条件下,随着氮添加水平升高,接合菌门相对丰度呈降低趋势,子囊菌门相对丰度变化则不明显。在相同的氮添加水平下,水分添加使接合菌门相对丰度增加,而担子菌门、子囊菌门、球囊菌门和壶菌门的相对丰度降低。在不同氮素和水分添加条件下,有5个土壤真菌门类11个真菌纲相对丰度变化显著。接合菌门的Mortierella属,担子菌门的Entolomataceae科和Geastrum属相对丰度变化极显著,可作为土壤真菌群落结构变化的指示种。PCo A分析结果也表明氮素和水分添加改变了土壤真菌群落结构。植物-土壤-微生物系统的结构方程模型结果表明,植物群落组成及植物物种丰富度的变化是土壤真菌群落结构发生变化的主要影响因素,土壤无机氮及p H的变化主要通过影响植物群落间接影响真菌群落,其对真菌群落的直接影响则较小。综上,氮素和水分添加改变了土壤真菌群落结构,且两者存在明显的互作效应,水分添加可改变氮添加对土壤真菌群落的影响。     

Carbon and Nitrogen Decoupling Under an 11-Year Drought in the Shortgrass Steppe

The frequency and magnitude of drought is expected to increase in the US Great Plains under future climate regimes. Although semiarid systems are considered highly resistant to water limitation, novel drought events could alter linkages among biogeochemical processes, and result in new feedbacks that influence the timescale of ecosystem recovery. We examined changes in carbon and nitrogen cycling in the last 2 years of an 11-year drought manipulation in the shortgrass steppe, and under the first 2 years of recovery from drought. We measured plant production, plant tissue chemistry, soil trace gas flux, and soil inorganic nitrogen dynamics to test the extent that this magnitude of drought altered carbon and nitrogen fluxes and how these changes affected post-drought dynamics. We found that soil inorganic nitrogen was up to five times higher under severe drought than under control conditions, but that this nitrogen may not have been accessible to plants and microbial communities during drought due to diffusion limitations. Drought plots had higher N₂O flux when they received equal rainfall pulses, showing that this accumulated N may be vulnerable to loss. In addition, plants in drought plots had higher tissue nitrogen for 2 years following drought. These results show that decadal-length droughts that may occur under future precipitation regimes are likely to alter ecosystem properties through interactions among precipitation, vegetation, and N cycling. Shifts in plant N, vulnerability of nitrogen to loss, and rainfall use efficiency that we observed are likely to affect the recovery time of semiarid systems subject to droughts of this magnitude.     

Do “hot moments” become hotter under climate change? Soil nitrogen dynamics from a climate manipulation experiment in a post-harvest forest

Whole-tree forest harvest can increase soil nitrous oxide (N₂O) effluxes and leaching of nitrogen (N) from soils. These altered N dynamics are often linked to harvesting effects on microclimate, suggesting that this “hot moment” for N cycling may become hotter with climate change. We hypothesized that increases in temperature and precipitation during this post-harvest period would increase availability of soil mineral N and soil-atmosphere N₂O efflux. To test this hypothesis we implemented a climate manipulation experiment after a forest harvest, and measured soil N₂O fluxes and inorganic N accumulating on ion exchange resins. Climate treatments were: control (A, ambient), heated (H, +2.5 °C), wetted (W, +23 % precipitation), and a two-factor treatment (H+W). For all treatments, the first year after harvest had highest N₂O efflux and resin N. Wetting significantly increased cumulative soil N₂O fluxes, but only when soils were not heated too. The cumulative soil-to-atmosphere N₂O efflux from W (5.8 mg N₂O–N m^?2) was significantly higher than A (?1.9 mg N₂O–N m^?2), but H+W (~0 mg N₂O–N m^?2) was similar to A. Regardless of wetting, heating increased resin N, but only on certain dates. Cumulative resin N was on average 125 % greater in the H plots than non-heated plots. Thus, changes in temperature and precipitation each impart distinct changes to the soil N cycle. Heating increased resin N regardless of water inputs, while wetting increasing N₂O but not when combined with heating. Our results suggest that climate change may exacerbate soil N losses from whole-tree harvest in the future, but the form and quantity of N loss will depend on how the future climate changes.     

氮添加对宁夏荒漠草原植物初级生产力的影响机制

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

Transpiration of a 64-year old maritime pine stand in Portugal

Alpine plant species have been shown to exhibit a more pronounced increase in leaf photosynthesis under elevated CO₂ than lowland plants. In order to test whether this higher carbon fixation efficiency will translate into increased biomass production under CO₂ enrichment we exposed plots of narrow alpine grassland (Swiss Central Alps, 2470 m) to ambient (355 μl l^-1) and elevated (680 μl l^-1) CO₂ concentration using open top chambers. Part of the plost received moderate mineral nutrient additions (40 kg ha^-1 year^-1 of nitrogen in a complete fertilizer mix). Under natural nutrient supply CO₂ enrichment had no effect on biomass production per unit land area during any of the three seasons studied so far. Correspondingly, the dominant species Carex curvula and Leontodon helveticus as well as Trifolium alpinum did not show a growth response either at the population level or at the shoot level. However, the subdominant generalistic species Poa alpina strongly increased shoot growth (+47%). Annual root production (in ingrowth cores) was significantly enhanced in C. curvula in the 2nd and 3rd year of investigation (+43%) but was not altered in the bulk samples for all species. Fertilizer addition generally stimulated above-ground (+48%) and below-ground (+26%) biomass production right from the beginning. Annual variations in weather conditions during summer also strongly influenced above-ground biomass production (19–27% more biomass in warm seasons compared to cool seasons). However, neither nutrient availability nor climate had a significant effect on the CO₂ response of the plants. Our results do not support the hypothesis that alpine plants, due to their higher carbon uptake efficiency, will increase biomass production under future atmospheric CO₂ enrichment, at least not in such late successional communities. However, as indicated by the response of P. alpina, species-specific responses occur which may lead to altered community structure and perhaps ecosystem functioning in the long-term. Our findings further suggest that possible climatic changes are likely to have a greater impact on plant growth in alpine environments than the direct stimulation of photosynthesis by CO₂. Counter-intuitively, our results suggest that even under moderate climate warming or enhanced atmospheric nitrogen deposition positive biomass responses to CO₂ enrichment of the currently dominating species are unlikely.     

Using plant functional traits to explain community composition across a strong environmental filter in Australian alpine snowpatches

Environmental filters act to limit the local community assemblage from the regional species pool by restricting the viable trait states that can occur there. In alpine snowpatches, the timing of snowmelt is a strong environmental filter. In coming decades, the strength of this filter is likely to relax with global climate change. We used three continuous plant functional traits (leaf area, plant height, seed mass) and their divergence (using the FD_var index) to document current patterns of community assembly and predict plant community responses to future environmental filters in alpine snowpatch vegetation. The community trait-weighted mean for leaf area and height, but not seed mass, was significantly higher in early snowmelt zones relative to mid and late melting zones across all snowpatches. Mean FD_var for height (but not leaf area or seed mass), by contrast, was substantially lower in early snowmelt zones, indicating that species growing in early melt zones are consistently taller than those growing in other zones. These results suggest that if climate change leads to earlier snowmelt and hence, a longer growing season, taller (more competitive) species with larger leaf areas (more productive) may replace short species in snowpatches as these plant communities re-assemble in response to changing environmental filters.     

温度增高、CO2浓度升高、施氮对蒙古栎幼苗非结构碳水化合物积累及其分配的综合影响

蒙古栎(Quercus mongolica)是中国东北地区天然次生林重要组成树种, 研究该树种幼苗有机碳积累及碳库容对未来气候变化的响应, 可为预测未来气候变暖情景下蒙古栎林的天然更新及幼苗的培育提供科学参考。该文旨在探讨CO₂浓度和温度升高综合作用对蒙古栎幼苗非结构性碳水化合物(NSC)积累及其分配的影响。实验环境条件用人工气候箱控制, 控制条件如下: 1) CO₂浓度倍增(700 μmol·mol^-1), 温度升高4 ℃处理(HCHT); 2) CO₂浓度正常(400 μmol·mol^-1), 温度升高4 ℃处理(HT); 3) CO₂浓度和温度均正常, 即对照组(CK); 每个气候箱幼苗分别在3种氮素水平下生长: N2 (15 mmol·L^-1, 高氮), N1 (7.5 mmol·L^-1, 正常供氮)和N0 (不施氮), 一共为9个处理。研究结果表明, 1) HCHT共同作用对NSC积累无促进作用, 但改变了植物各器官中NSC的分配比例, 叶片中可溶性糖和淀粉的积累明显增加, HCHT下N2水平有利于NSC的积累。2) HT明显影响了蒙古栎一年生幼苗NCS的积累和分配。在N2水平下, HT明显促进NSC的积累, 并增加了在主根中的分配比例。3)植株各器官可溶性糖含量的动态变化因处理不同而异。主根淀粉含量随时间逐渐增加, 而细根淀粉含量随时间逐渐减少。在未来气候变暖的情况下, 土壤中大量的氮供给, 可能将促进蒙古栎幼苗的生长、增加其碳库容和抵御不良环境的能力, 进而提高其天然更新潜力。     

Climate change drives a shift in peatland ecosystem plant community: Implications for ecosystem function and stability

The composition of a peatland plant community has considerable effect on a range of ecosystem functions. Peatland plant community structure is predicted to change under future climate change, making the quantification of the direction and magnitude of this change a research priority. We subjected intact, replicated vegetated poor fen peat monoliths to elevated temperatures, increased atmospheric carbon dioxide (CO₂), and two water table levels in a factorial design to determine the individual and synergistic effects of climate change factors on the poor fen plant community composition. We identify three indicators of a regime shift occurring in our experimental poor fen system under climate change: nonlinear decline of Sphagnum at temperatures 8 °C above ambient conditions, concomitant increases in Carex spp. at temperatures 4 °C above ambient conditions suggesting a weakening of Sphagnum feedbacks on peat accumulation, and increased variance of the plant community composition and pore water pH through time. A temperature increase of +4 °C appeared to be a threshold for increased vascular plant abundance; however the magnitude of change was species dependent. Elevated temperature combined with elevated CO₂ had a synergistic effect on large graminoid species abundance, with a 15 times increase as compared to control conditions. Community analyses suggested that the balance between dominant plant species was tipped from Sphagnum to a graminoid‐dominated system by the combination of climate change factors. Our findings indicate that changes in peatland plant community composition are likely under future climate change conditions, with a demonstrated shift toward a dominance of graminoid species in poor fens.     

Plant species richness, elevated CO2, and atmospheric nitrogen deposition alter soil microbial community composition and function

We determined soil microbial community composition and function in a field experiment in which plant communities of increasing species richness were exposed to factorial elevated CO₂ and nitrogen (N) deposition treatments. Because elevated CO₂ and N deposition increased plant productivity to a greater extent in more diverse plant assemblages, it is plausible that heterotrophic microbial communities would experience greater substrate availability, potentially increasing microbial activity, and accelerating soil carbon (C) and N cycling. We, therefore, hypothesized that the response of microbial communities to elevated CO₂ and N deposition is contingent on the species richness of plant communities. Microbial community composition was determined by phospholipid fatty acid analysis, and function was measured using the activity of key extracellular enzymes involved in litter decomposition. Higher plant species richness, as a main effect, fostered greater microbial biomass, cellulolytic and chitinolytic capacity, as well as the abundance of saprophytic and arbuscular mycorrhizal (AM) fungi. Moreover, the effect of plant species richness on microbial communities was significantly modified by elevated CO₂ and N deposition. For instance, microbial biomass and fungal abundance increased with greater species richness, but only under combinations of elevated CO₂ and ambient N, or ambient CO₂ and N deposition. Cellobiohydrolase activity increased with higher plant species richness, and this trend was amplified by elevated CO₂. In most cases, the effect of plant species richness remained significant even after accounting for the influence of plant biomass. Taken together, our results demonstrate that plant species richness can directly regulate microbial activity and community composition, and that plant species richness is a significant determinant of microbial response to elevated CO₂ and N deposition. The strong positive effect of plant species richness on cellulolytic capacity and microbial biomass indicate that the rates of soil C cycling may decline with decreasing plant species richness.     

Loss of plant biodiversity eliminates stimulatory effect of elevated CO2 on earthworm activity in grasslands

Earthworms are among the world’s most important ecosystem engineers because of their effects on soil fertility and plant productivity. Their dependence on plants for carbon, however, means that any changes in plant community structure or function caused by rising atmospheric CO₂ or loss of plant species diversity could affect earthworm activity, which may feed back on plant communities. Production of surface casts measured during three consecutive years in field experimental plots (n = 24, 1.2 m²) planted with local calcareous grassland species that varied in plant species richness (diversity levels: high, 31 species; medium, 12; low, 5) and were exposed to ambient (356 μl CO₂ l^?1) or elevated (600 μl CO₂ l^?1) CO₂ was only consistently stimulated in high diversity plots exposed to elevated CO₂ (+120 %, 31 spp: 603 ± 52 under ambient CO₂ vs. 1,325 ± 204 g cast dwt. m^?2 year^?1 under elevated CO₂ in 1996; +77 %, 940 ± 44 vs. 1,663 ± 204 g cast dwt. m^?2 year^?1 in 1998). Reductions in plant diversity had little effect on cast production in ecosystems maintained at ambient CO₂, but the stimulatory effect of elevated CO₂ on cast production disappeared when plant species diversity was decreased to 12 and 5 species. High diversity plots were also the only communities that included plant species that an earlier field study showed to be among the most responsive to elevated CO₂ and to be most preferred by earthworms to deposit casts near. Further, the +87 % CO₂-induced increase in cast production measured over the 3 years corresponded to a parallel increase in cumulative total nitrogen of 5.7 g N m^?2 and would help explain the large stimulation of aboveground plant biomass production observed in high-diversity communities under elevated CO₂. The results of this study demonstrate how the loss of plant species from communities can alter responses of major soil heterotrophs and consequently ecosystem biogeochemistry.     

Ectomycorrhizal fungal succession coincides with shifts in organic nitrogen availability and canopy closure in post-wildfire jack pine forests

Successional changes in belowground ectomycorrhizal fungal (EMF) communities have been observed with increasing forest stand age; however, mechanisms behind this change remain unclear. It has been hypothesized that declines of inorganic nitrogen (N) and increases of organic N influence changes in EMF taxa over forest development. In a post-wildfire chronosequence of six jack pine (Pinus banksiana) stands ranging in age from 5 to 56 years, we investigated EMF community composition and compared shifts in taxa with detailed soluble inorganic and organic N data. Taxa were identified by internal transcribed spacer rDNA sequencing, and changes in community composition evaluated with non-metric multi-dimensional scaling (NMDS). Dissimilarities in the community data were tested for correlations with N variables. We observed a successional shift along NMDS axis 1 from such taxa as Suillus brevipes and Thelephora terrestris in sites age 5 and 11 to species of Cortinarius and Russula, among others, in the four older sites. This change was positively correlated with soluble organic N (SON) (r ² = 0.902, P = 0.033) and free amino-acid N (r ² = 0.945, P = 0.021), but not inorganic N. Overall, our results show a successional shift of EMF communities occurring between stand initiation and canopy closure without a change in species of the dominant plant–host, and associated with SON and free amino-acid N in soil. It is uncertain whether EMF taxa are responding to these organic N forms directly, affecting their availability, or are ultimately responding to changes in other site variables, such as belowground productivity.     

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

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

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

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

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

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