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Forest productivity and carbon (C) sequestration largely depend on soil N and P availability. To date, however, the temporal variation of nutrient limitation along forest succession is still under debate. Leaf stoichiometry and nutrient resorption are important indicators for predicting nutrient limitation of plant growth. Here, we measured nitrogen (N) and phosphorus (P) concentrations in green leaves and leaf litter for all woody species at four stages of temperate forest succession, and analyzed how abiotic and biotic factors affect leaf stoichiometry and nutrient resorption along forest succession. At the individual scale, leaf N and P concentrations had a significant increase at the end of the succession, while no change in leaf N:P ratio was detected. Nitrogen resorption efficiency (NRE) increased significantly with succession, but P resorption efficiency (PRE) first increased and then decreased. Significant increases in NRE:PRE ratios only occurred at the end of the succession. Moreover, plant N cycling was less responsive to soil nutrient than P cycling. At the community scale, we found that leaf N and P concentrations first decreased and then increased along forest succession, which were mainly affected by Shannon–Wiener index and species richness. Leaf N:P ratio significantly varied with succession and was mainly determined by community-weighted mean diameter at breast height (DBH). NRE increased and was significantly influenced by species richness and DBH, while PRE was relatively stable along forest succession. Thus, the NRE:PRE ratios significantly increased, indicating that N limitation is exacerbated with the temperate forest succession. These results might reflect the intense interspecific competition for limiting resource in a higher biodiversity community. In conclusion, our findings highlight the importance of biotic factors in driving forest ecosystem nutrient cycling and provide valuable information for sustainable fertilizer management practices in China’s temperate and boreal forests.

Elucidating mechanisms underlying community assembly and biodiversity patterns is central to ecology and evolution. Genome size (GS) has long been hypothesized to potentially affect species' capacity to tolerate environmental stress and might therefore help drive community assembly. However, its role in driving β-diversity (i.e., spatial variability in species composition) remains unclear. We measured GS for 161 plant species and community composition across 52 sites spanning a 3200-km transect in the temperate grasslands of China. By correlating the turnover of species composition with environmental dissimilarity, we found that resource filtering (i.e., environmental dissimilarity that includes precipitation, soil nitrogen and phosphorus concentrations) affected β-diversity patterns of large-GS species more than small-GS species. By contrast, geographical distance explained more variation of β-diversity for small-GS than for large-GS species. In a 10-year experiment manipulating levels of water, nitrogen, and phosphorus, adding resources increased plant biomass in species with large-GS, suggesting that large-GS species are more sensitive to the changes of resource availability. These findings highlight the role of GS in driving community assembly and predicting species responses to global change.

Soil nematodes play a fundamental role in regulating ecosystem carbon and nutrient cycling. It is widely recognized that soil nematode community composition is sensitive to nutrient enrichment, but the linkage between community assembly processes and functional changes under nutrient enrichment condition remains poorly understood. We examined the compositional and functional responses and quantified the role of main community assembly processes (genus losses, genus gains, and context-dependent variations of abundance) in driving the carbon budget of soil nematode communities in response to nitrogen (N) and phosphorus (P) addition in a temperate grassland. Nitrogen and P addition significantly interacted to affect nematodes abundance, biomass, and functional variables of C cycling, in that P addition increased all the variables under ambient N condition but not under N enriched condition. Soil pH, ammonium concentration, and total phosphorus concentration played important roles in driving the variations of nematode C budgets, indicating the minor role of plant community characteristics. The enhancement of all variables following P addition was caused by the increases in the abundance of common genera (e.g. Acrobeles, Scutylenchus, and Tylencholaimus). The variation of genus richness contributed to the P-induced increases of nematode abundance but not to the increases of carbon budgets. Our results uncover the linkages between community assembly processes and the abundance and C cycling function of soil nematode community under nutrient enrichment conditions. The significant interactive effects between N and P addition highlight the complexity in predicting the compositional and functional changes in soil nematode community under a scenario of multiple-nutrient enrichment.

The concentrations and stoichiometry of soil carbon (C), nitrogen (N), and phosphorus (P) have critical implications for nutrient cycling and ecosystem function. While their high sensitivity to atmospheric N deposition is well known, it remains unclear for the soil depth-dependence of such responses to N deposition. Here, we examined the responses of soil C : N : P stoichiometry at three soil depths in the upper humus horizon (0–5, 5–10, and 10–20 cm) of Haplic Chernozem (Loamic) across a gradient of urea addition rates (0, 2, 10, 20, and 50 g N m–2 year–1) after five years treatments in a hay-harvest meadow steppe of northern China. We found that the effects of increasing N addition rates on the concentrations and stoichiometry of soil C, N and P did not depend on soil depth, though those parameters varied greatly across different soil layers. Across all soil depths, the concentrations of soil C and N increased with increasing N addition rates, but soil P concentration was not affected by N addition. The higher sensitivity of soil N than soil C to N enrichment resulted in decreasing soil C : N ratio across the N addition gradient, especially for the surface soil layer. Soil N : P ratio showed a positive response to the increases of N addition rates. The unbalanced responses of soil C, N, and P concentrations to N enrichment, as indicated by the decreases of soil C : N ratio and the increases of soil N : P ratio across the N addition gradient, highlighted the role of N enrichment in decoupling soil nutrient cycling in the temperate steppe.

Calcium (Ca) is an essential nutrient for plant growth and Ca concentrations in forage have important implications for ruminant diet and health. It remains an open question whether forage Ca concentration would be decreased by increasing nitrogen (N) deposition. We manipulated the increasing rates of N addition (2008–2015) in a semiarid grassland, northern China. Plant Ca concentrations for all species were examined in each plot under N treatment. The Ca concentrations at functional group and community levels were calculated based on the concentration of each species presented and their relative biomass in each plot. We found that community-level Ca concentration remained stable across a gradient of wide-ranged N addition rates, although Ca concentration at both species and functional group levels showed negative responses to N enrichment. Given that forbs had higher Ca concentration than grasses, the increasing relative biomass of forbs canceled out the negative responses of species-level and functional group-level Ca concentration. Our results further showed that community Ca pool showed a positive but saturating response to N addition, with a threshold at the rate of 10 g N m−2 yr−1. Our findings highlight the role of changes in plant relative biomass in controlling the responses of forage Ca concentration and stock to N enrichment.

Aims Plant nutrient use efficiency, an important factor driving primary production, is sensitive to nitrogen (N) deposition. Because of strengthened regulations of N emissions, atmospheric N deposition is decreasing in many countries. It remains unknown whether historical N enrichment would alter plant nutrient use efficiency and whether such impacts would be mitigated by ecosystem management strategy. Methods We assessed the effects of historical N addition and mowing on plant N and phosphorus use efficiency (NUE and PUE) at both functional group and community levels in a temperate steppe after the cessation of 6-year N addition. Important Findings Historical N addition had negative legacy effects on plant NUE but not on PUE at the functional group level. There were negative legacy effects of N addition on community-level NUE and PUE, but only in the unmown plots. Mowing mitigated the negative legacy effects of N enrichment on community-level NUE and PUE by reducing the dominance of tall rhizomatous grass but enhancing that of tall bunchgrass. Our results highlight the importance of community composition variations caused by mowing in driving the legacy effects of N enrichment on community-level nutrient use efficiency. Given the slow recovery of community composition following cessation of N deposition, our findings suggest that the lower nutrient efficiency of plant community would be long-lasting.

Background Forage production is the fundamental ecosystem service of grasslands. Although forage consumption occurs at community level, most studies focused on species-level changes of forage quality. The quantitative and qualitative changes of forage production are driven by species-specific trait, intra-specific plasticity, and species turnover. We examined the changes in forage production and digestibility after 5-year factorial treatments of nitrogen (N) addition and mowing in a temperate steppe and linked such changes to community assembly under the Price equation framework. Results Nitrogen addition significantly reduced species richness, increased forage production, but did not change forage digestibility (indicated by the total Ca+Mg concentrations). Mowing did not affect forage production and digestibility. The positive effects of N addition on forage production were driven by the enhancement of abundance of the remaining species following N enrichment, rather than by species loss or species gain. The species identity effects could offset the effects of species richness loss or gain on forage production and digestibility. Conclusions Our results highlight the importance of a community perspective in addressing the quantitative and qualitative changes of forage production under global change pressure of N enrichment. Species identity is important in determining the contribution of different processes of community assembly to ecosystem services.

Aims Understanding the patterns and drivers of carbon isotope discrimination (13Δ) in C3 and C4 functional groups is critical for predicting C3/C4 vegetation ratio from the isotopic composition of soil organic matter. In this study, we aimed to evaluate how intraspecific variation will modify functional group-level 13Δ values and the associated prediction of C3/C4 vegetation ratio. Methods We investigated 13Δ of 726 individual plants (96 species; C3 and C4 functional groups) and topsoil organic matter in 26 grassland communities along an aridity gradient in northern China. The fraction of C4 contribution was calculated with mixing models that considered: (i) both intra- and interspecific effects on the 13Δ values of C3 and C4 functional groups; (ii) only interspecific effects; or (iii) none of these effects. Important Findings We found divergent responses of plant 13Δ at the intraspecific level to the changes of aridity across the gradient. The 13Δ of both C3 and C4 functional groups was negatively correlated with an aridity index, with higher sensitivity for C3 than for C4 functional groups. Intraspecific 13Δ variation played a key role in driving the total 13Δ variations of C3 plants. Overlooking such intraspecific effect in mixing models led to a greatly increased fraction of C4 contribution to soil organic carbon. A correction for the effects of intraspecific variation is therefore essential for correctly inferring C3/C4 vegetation ratio in the past. Our findings provide basic information for the reconstruction of past vegetation change from bulk materials in arid and semiarid biomes.

Non-additive effects during the decomposition of mixed litter at species level have important consequences on ecosystem nutrient cycling, whereas such effect at plant organ level remains unclear. We investigated mass loss and nutrient release of single and mixed litter from leaf and culm for a dominant grass and of shoots for two dominant grasses under both ambient and enriched N conditions in a temperate grassland. We found comparable mixing effects on litter mass loss and nutrient release at organ and species levels after 2-yr decomposition. Nitrogen enrichment stimulated litter mass loss of all litter types but did not alter the mixing effects on mass loss. Further, N enrichment enhanced the non-additive effects of mixing on N release at plant organ level but not at species level. This study extend the non-additive effects of litter decomposition from inter-specific level to intra-specific level by highlighting the synergistic interaction between leaf litter and culm litter during decomposition. Given the existence of non-additive mixing effects at plant organ level, it is more difficult to predict litter decomposition and nutrient cycling in herbaceous communities.

The continuing nitrogen (N) deposition observed worldwide alters ecosystem nutrient cycling and ecosystem functioning. Litter decomposition is a key process contributing to these changes, but the numerous mechanisms for altered decomposition remain poorly identified. We assessed these different mechanisms with a decomposition experiment using litter from four abundant species (Achnatherum sibiricum, Agropyron cristatum, Leymus chinensis and Stipa grandis) and litter mixtures representing treatment-specific community composition in a semi-arid grassland under long-term simulation of six different rates of N deposition. Decomposition increased consistently with increasing rates of N addition in all litter types. Higher soil manganese (Mn) availability, which apparently was a consequence of N addition-induced lower soil pH, was the most important factor for faster decomposition. Soil C : N ratios were lower with N addition that subsequently led to markedly higher bacterial to fungal ratios, which also stimulated litter decomposition. Several factors contributed jointly to higher rates of litter decomposition in response to N deposition. Shifts in plant species composition and litter quality played a minor role compared to N-driven reductions in soil pH and C : N, which increased soil Mn availability and altered microbial community structure. The soil-driven effect on decomposition reported here may have long-lasting impacts on nutrient cycling, soil organic matter dynamics and ecosystem functioning.

Nutrient addition is a widely-used strategy to restore degraded grasslands. It remains unknown whether and how the number of added nutrients affects the soil nematode community in degraded grasslands. We examined the immediate responses of taxonomic and functional composition of the soil nematode community to different numbers of added nutrients using factorial combinations of nitrogen (N), phosphorus (P), and potassium (K + micronutrients) in a degraded grassland of northern China. The taxonomic and functional composition of the soil nematode community generally formed a unimodal relationship with the number of added nutrients. Changes of soil pH after fertilization affected the structural stability and complexity of the soil nematode community. The magnitude of nematode functional responses to nutrient supply was driven by changes of plant aboveground biomass and soil pH. Soil nematode community showed non-linear responses to the variations of the number of added nutrients in a degraded grassland, which contrasts previous findings from plant community. The number of added nutrients should be given full consideration in formulating effective restoration strategies for degraded grasslands.

Anthropogenic nitrogen (N) deposition has affected plant community composition and nutrient cycling in terrestrial ecosystems worldwide. This includes changes to the way plants use and recycle nutrients, including effects on nutrient resorption, which is a key process through which plants recover nutrients from tissue during senescence. Nutrient resorption has considerable adaptive and functional significance for plants and helps regulate core ecosystem processes such as decomposition. However, our understanding of how N deposition affects nutrient resorption and, in particular, of how N inputs alter ecosystem resorption via changes to existing species’ resorption compared with changes to community composition remains poor. To disentangle the role of species versus community composition controls driving variation in nutrient resorption responses to N inputs, we carried out an experiment with six different N addition rates in a temperate steppe. We found that species-scale nutrient resorption responses to N enrichment were variable; for example, only half of the measured species reduced both N and P resorption efficiency in response to increased N inputs. In contrast, community-scale responses consistently resulted in reduced N and P resorption. Still, N-induced changes in community composition were a weaker control on overall resorption responses than were the effects on individual species; however, it was the synergistic interaction between the two that resulted in the large total reductions of nutrient resorption in the face of increased N. Taken together, our results highlight that understanding and predicting nutrient resorption responses will be most accurately scaled by accounting not only for species’ reductions in resorption but also for changes in community composition.

Uncovering the importance of soil and plant characteristics in driving the legacy effects of nitrogen (N) deposition on plant community nutrient stoichiometry would improve our understanding of plant-soil interaction during restoration of historically N-enriched ecosystems. Based on a field experiment with the cessation of six-year N addition in a temperate steppe of northern China, we measured concentrations and stoichiometry of N and phosphorus (P) in soils and different plant functional groups, under both mown and unmown conditions. Historical N addition did not affect soil total and available N and P concentrations and stoichiometry, but significantly altered plant community composition. Plant nutrient concentrations and N:P ratios significantly differed among four plant functional groups. The concentrations and stoichiometric ratios of N and P between soils and plants were generally not correlated. The positive legacy effects of N addition on community N:P stoichiometry were caused by the biomass enhancement of tall bunchgrass, the functional group with the highest N:P ratios. Changes in plant community composition instead of soil nutrient status were the main driver for the positive legacy effects of N enrichment on plant community stoichiometry. Given that the recovery of community composition after the cessation of N deposition is generally slow, our findings indicate that the legacy effects of N deposition on soil nutrient cycling would persist in long-term due to the importance of plant-mediated pathway.

The stoichiometric traits of litter play an important role in driving litter decomposition and ecosystem nutrient cycling. While the impacts of nitrogen (N) deposition on the species-level litter stoichiometric traits have been well addressed, we know little about that at community-level, which is supposed to be driven by both intra-specific variation and changes in community composition. We examined the effects of N deposition on litter phosphorus (P) concentration and N:P ratio at both species- and community-level in a semi-arid grassland of northern China. We further decomposed the community-level variations of litter nutritional traits into intra- and inter-specific variation. Nitrogen addition, especially at high rates, substantially changed community composition. Litter P concentrations and N:P ratios significantly varied among different species. Litter P concentrations and N:P ratios at both species- and community-level were positively correlated with N addition rates. Biomass-weighted community-level N:P ratios were more sensitive to N addition than the non-weighted ones, indicating that community composition strengthened the positive impacts of N addition on litter N:P ratios. There was positive co-variation between intra- and inter-specific variation for litter N:P ratio, indicating the consistency of community composition and intra-specific variation in their effects on litter N:P ratio. Our results indicated that the imbalance of N and P following N enrichment would be much larger than the expectation based on the findings from species-level, and thus highlight the importance of changes in community composition in driving the responses of community-level litter N:P stoichiometry to N deposition in the semi-arid grassland.

Human-induced nitrogen (N) and phosphorus (P) enrichment have profound effects on grassland net primary production (NPP) and species richness. However, a comprehensive understanding of the relative contribution of N vs. P addition and their interaction on grassland NPP increase and species loss remains elusive. We compiled data from 80 field manipulative studies and conducted a meta-analysis (2107 observations world-wide) to evaluate the individual and combined effects of N and P addition on grassland NPP and species richness. We found that both N addition and P addition significantly enhanced grassland above-ground NPP (ANPP; 33.2% and 14.2%, respectively), but did not affect total NPP, below-ground NPP (BNPP), and species evenness. Species richness significantly decreased with N addition (11.7%; by decreasing forbs) probably due to strong decreased soil pH, but not with P addition. The combined effects of N and P addition were generally stronger than the individual effects of N or P addition, and we found the synergistic effects on ANPP, and additive effects on total NPP, BNPP, species richness, and evenness within the combinations of N and P addition. In addition, N and P addition effects were strongly affected by moderator variables (e.g. climate and fertilization type, duration and amount of fertilizer addition). These results demonstrate a higher relative contribution of N than P addition to grassland NPP increase and species loss, although the effects varied across climate and fertilization types. The existing data also reveals that more long-term (≥5 years) experimental studies that combine N and P and test multifactor effects in different climate zones (particularly in boreal grasslands) are needed to provide a more solid basis for forecasting grassland community response and C sequestration response to nutrient enrichment at the global scale.

The nitrogen‑phosphorus (N-P) imbalance induced by N enrichment has received increasing concerns, because N:P ratios play a critical role in driving many fundamental ecological processes. Given the simultaneous occurrence of different global change drivers, it is important to understand whether and how would such N-induced N-P imbalance would be mediated by other global change factors. We examined the interactive effects of N addition (10 g N m

Nitrogen (N) deposition has negative consequences for plant communities and ecosystem functioning. Nitrogen deposition is declining or is projected to decline in many regions due to the reduction of N-compound emissions. It remains unknown whether the cessation of N deposition will have legacy effects on plant nutrient stoichiometry. If there are stoichiometric legacy effects at the community-level from prior N enrichment, the contribution of changes in community composition relative to the stoichiometric responses of individual functional groups in driving these changes is unknown. We measured plant N:P ratio at both functional group level and community level in a temperate steppe in three consecutive years following cessation of a 7-year N addition experiment with a wide-range of total amount added N in which the total amount of N added ranged from 0 to 350 g m−2. The total amount of N addition was positively correlated with tall rhizomatous grass productivity and negatively correlated with tall bunchgrass productivity. Plant N:P ratios at both functional group and community levels did not change across N addition gradient. Community composition shifts contributed less than intra-group variation to the total variation of community level nutrient status in response to historic N addition. Our results indicated that biological N:P imbalance in response to N deposition can be rapidly ameliorated after the cessation of N deposition. Such legacy effects of N deposition on plant nutrient status are largely driven by the changes of nutrient status of individual plants instead of N-induced shifts in community composition.

Atmospheric nitrogen (N) deposition, generally, has been simulated through a single or relatively few N applications per year for its ecological effect study. Despite the importance of timing in ecosystem processes, ecological experiments with more realistic N addition frequencies are rare. We employed a novel design with typical twice (2X) versus atypical monthly (12X) N applications per year to explore effects of N addition frequency on above‐ and below‐ground biodiversity and function. Each year, several response variables from either below‐ground or above‐ground growth, N status and cycling, or plant and bacterial diversity differed as a result of N addition frequency. BNPP showed a large frequency effect in the relatively moist year but not in the dry year. Nitrogen addition decreased root growth in the monthly relative to the biannual applications, which could be highly consequential for predicting changes in global carbon and nitrogen cycling. Simulated N deposition tended to perturb biodiversity, but it is noteworthy that 12X applications that spread N deposition more evenly through a year have much less negative impacts on plant and bacterial diversities than 2X amendments per year. Soil N mineralization rate in year 6 was much lower when N additions were monthly compared with a biannual amendment, especially when simulated N deposition was high. We have established that amendment frequency matters for understanding ecosystem response to N deposition. Experiments that more closely mimic the anthropogenic process of N deposition are needed to best assess ecosystem and potential global biogeochemical changes. A free Plain Language Summary can be found within the Supporting Information of this article.

Owing to different soil environments and organic carbon sources, soil organic carbon (SOC) composition and preservation mechanisms may vary substantially in deep versus surface soils. This study examines vertical variation of various SOC components (including lignin phenols, cutin, suberin and microbial lipids) across the Chinese-Mongolian grasslands. Lignin phenols and hydrolysable lipids were isolated by cupric oxide oxidation and alkaline hydrolysis, respectively. A comprehensive list of environmental variables was compiled to disentangle influencing factors for the variation of various components at four different depths. Lignin phenols, suberin and microbial lipids were most abundant in the surface soil (0–10 cm). Ratios of cutin to suberin and microbial lipids to lignin phenols (in the alpine grasslands) were higher in the bottom-most soil. Calcium (Ca) rather than iron (Fe) plays an important role in the preservation of cutin and microbial lipids in the bottom soil with a high Ca:Fe ratio and high pH. The incorporation efficiency of suberin was higher in deeper than surface soils and showed an increasing trend with depth in the alpine (but not temperate) grasslands. Compared to surface soils, the incorporation of root-derived carbon into SOC is more efficient in deeper soils due to stronger mineral protection and weaker microbial decomposition. Root-derived carbon is more important for SOC accrual in the alpine than temperate grasslands, especially at depth. These findings reveal differential distribution patterns and preservation mechanisms for SOC components in the surface versus deep soils, providing new information to understand SOC stability at different depths.

Variations in plant nitrogen (N) and phosphorus (P) concentrations and ratios have great implications for primary productivity and nutrient cycling. Here, we reported their changes at functional group and community levels along an aridity gradient. We carried out a field investigation and set up 26 sampling sites along a transect in the drylands of northern China to examine the responses of plant nutrient status to increasing aridity and the relationships between plant and soil nutrients. The responses of N and P concentrations to increasing aridity differed between woody (sub-) shrubs and herbaceous species (including annual/biennial forbs and grasses, perennial forbs and perennial grasses). Nitrogen and P concentrations of shrubs decreased while those of herbs generally increased with increasing aridity. Plant N:P ratios increased significantly with increasing aridity for the plant functional groups and at the community level. At both organizational levels, nutrient concentrations in plants were negatively related to those in soils. Our results indicated that the responses of plant nutrition to aridity were dependent on plant functional group. Plant growth appeared increasingly limited by P, and plant and soil nutrients become increasingly decoupled at large spatial scale with increasing aridity.

Aims The stoichiometric characteristics of plant communities are important controller for several fundamental ecological processes. The effects of environmental changes on community stoichiometric characteristics are driven by intra- and inter-specific variation. However, the relative importance of both pathways has seldom been empirically examined. Methods We quantified the relative contribution of intra- and inter-specific variation to the changes of community nitrogen (N) and phosphorus (P) concentrations after seven-year factorial N addition and mowing treatments in a semi-arid grassland of northern China. Results Nitrogen addition significantly increased community N and P concentrations and N:P ratio. Mowing significantly increased community N concentration and N:P. Intra-specific variation contributed more than inter-specific variation to the total variability of all the nutritional and stoichiometric characteristics, with intra-specific variation accounting for 68%, 70%, and 75% of the total variation in community-level N, P, and N:P, respectively. Negative covariations between the contribution of intra- and inter-specific variation occurred for community N and P concentrations. Further, N addition and mowing interacted to affect the impacts of intra- and inter-specific variation on community N concentration and N:P stoichiometry. Conclusions Our results highlight different ways of trait selection for N addition and mowing treatments. Interactions between those two factors make it more difficult to accurately predict the responses of plant-mediated biogeochemical cycles under co-occurrence of environmental changes.

Between‐species variation in nutrient resorption is one of the mechanisms explaining the positive relationship between biodiversity and primary productivity. Yet, the role of within‐species variations in nutrient resorption in mediating the relationship between biodiversity and productivity remains unclear. We examined how within‐species nutrient resorption, and ultimately productivity, respond to changes in species richness by using four traits related to nitrogen and phosphorus use in four dominant species from different plant functional groups in a biodiversity removal experiment in the temperate steppe. Nitrogen and phosphorus concentrations in both green and senesced leaves in all species significantly decreased with increasing plant species richness, suggesting that plants used those limiting nutrients more efficiently with increasing biodiversity. Plants in higher diversity communities resorbed more nutrients during senescence, which may facilitate reproduction and vegetative regrowth in the next year. Synthesis. Our results highlight the importance of considering within‐species variation in nutrient resorption as an important underlying mechanism explaining the positive effects of biodiversity on primary productivity and ecosystem carbon accumulation.

As a major plant-derived soil organic carbon (SOC) component, lignin-derived phenolic compounds show varying biogeochemical characteristics compared to plant-derived lipid moieties. Comparing their distribution patterns can provide information on mechanisms governing SOC preservation and dynamics. However, the large-scale distribution pattern and stability of lignin versus plant-derived lipids are still poorly constrained. Here we investigated the distribution of lignin phenols versus plant-derived lipids in the surface soils across the alpine versus temperate grasslands of China and Mongolia. Lignin phenols were isolated by cupric oxide oxidation method and compared with the previously analyzed plant-derived lipids (cutin and suberin). A comprehensive list of environmental variables was compiled to disentangle the climatic, edaphic and vegetation influences on lignin phenols’ distribution in the soil. Lignin phenols showed similar SOC-normalized concentrations in the alpine and temperate grassland soils despite a higher plant input to the latter, suggesting better lignin preservation in the cold region. However, compared with plant-derived lipids (cutin and suberin), lignin seems to be less stabilized. The variation of lipid versus lignin components is mainly related to climate (particularly aridity) in the alpine grassland soils, while the relative abundance of plant lipids and lignin phenols is more related to reactive mineral contents in the temperate grassland soils. Lignin contributes differentially to SOC accumulation in the alpine and temperate soils: while lignin seems to be better preserved in the cold region, lignin phenols decrease relative to other carbon components with SOC accrual in the temperate region. Overall, lignin distribution and fate may be more sensitive to carbon source variations than temperature shifts in the grasslands.

Increasing availability of reactive nitrogen (N) threatens plant diversity in diverse ecosystems. While there is mounting evidence for the negative impacts of N deposition on one component of diversity, species richness, we know little about its effects on another one, species evenness. It is suspected that ecosystem management practice that removes nitrogen from the ecosystem, such as hay-harvesting by mowing in grasslands, would mitigate the negative impacts of N deposition on plant diversity. However, empirical evidence is scarce. Here, we reported the main and interactive effects of N deposition and mowing on plant diversity in a temperate meadow steppe with 4-year data from a field experiment within which multi-level N addition rates and multiple N compounds are considered. Across all the types of N compounds, species richness and evenness significantly decreased with the increases of N addition rate, which was mainly caused by the growth of a tall rhizomatous grass, Leymus chinensis. Such negative impacts of N addition were accumulating with time. Mowing significantly reduced the dominance of L. chinensis, and mitigated the negative impacts of N deposition on species evenness. We present robust evidence that N deposition threatened biodiversity by reducing both species richness and evenness, a process which could be alleviated by mowing. Our results highlight the changes of species evenness in driving the negative impacts of N deposition on plant diversity and the role of mowing in mediating such negative impacts of N deposition.