Soil extracellular enzyme activity (EEA) and enzyme stoichiometry are indicators of soil nutrient availability and microbial substrate limitation. However, the patterns in soil enzyme activities and stoichiometry, and their key drivers remain unclear during vegetation restoration in degraded sandy land. Here, soil ecosystems at five restoration stages of degraded sandy land were selected for investigation and we quantified the microbial nutrient limitation using EEA stoichiometry. The average C:N:P ratio for the nutrient-acquisition enzymes was 1:0.88:0.86, which differed from the global ratio of 1:1:1, indicating stronger microbial C limitation than N or P limitation. With vegetation restoration, the enzyme N:P ratio decreased, but the angle term used in vector analysis increased, indicating that the restored grassland transitioned from N-restricted (angle < 45°) to P-restricted (angle > 45°). Plant C inputs increased the soil nutrient content and significantly decreased the microbial C and N limitation but increased P limitation due to nutrient competition between plants and soil microorganisms. Decreased soil water levels caused by vegetation transpiration may have shifted the microbial limitation from N to P. The decreased fungi:bacteria ratio strengthened the microbial C and N or P limitations. On the basis of these findings, as measures to alleviate the associated nutrient limitations, we would recommend supplementation with phosphorus in the potential and slight stages of rocky desertification and the supplementary application of nitrogen in the moderate and severe stages during the restoration of degraded sandy ecosystems.

The decomposition of plant material in arid ecosystems is considered to be substantially controlled by water and N availability. The responses of litter decomposition to external N and water, however, remain controversial, and the interactive effects of supplementary N and water also have been largely unexamined. A 3.5-year field experiment with supplementary nitrogen and water was conducted to assess the effects of N and water addition on mass loss and nitrogen release in leaves and fine roots of three dominant plant species (i.e., Artemisia halondendron, Setaria viridis, and Phragmites australis) with contrasting substrate chemistry (e.g. N concentration, lignin content in this study) in a desertified dune grassland of Inner Mongolia, China. The treatments included N addition, water addition, combination of N and water, and an untreated control. The decomposition rate in both leaves and roots was related to the initial litter N and lignin concentrations of the three species. However, litter quality did not explain the slower mass loss in roots than in leaves in the present study, and thus warrant further research. Nitrogen addition, either alone or in combination with water, significantly inhibited dry mass loss and N release in the leaves and roots of the three species, whereas water input had little effect on the decomposition of leaf litter and fine roots, suggesting that there was no interactive effect of supplementary N and water on litter decomposition in this system. Furthermore, our results clearly indicate that the inhibitory effects of external N on dry mass loss and nitrogen release are relatively strong in high-lignin litter compared with low-lignin litter. These findings suggest that increasing precipitation hardly facilitates ecosystem carbon turnover but atmospheric N deposition can enhance carbon sequestration and nitrogen retention in desertified dune grasslands of northern China. Additionally, litter quality of plant species should be considered when modelling the carbon cycle and nutrient dynamics of this system.

The number of livestock per unit area is commonly used as a proxy of grazing pressure in both experimental studies and grassland management. However, this practice ignores the impact of landform heterogeneity on the spatial distribution of grazing pressure, leading to localized patches of degraded grassland. The spatial distribution of actual grazing density thus needs to be examined. Owing to the corresponding changes in resource availability and energy consumption as livestock move across an elevation gradient, we predict that livestock will preferentially use low-land and that different temporal patterns of grazing pressure will occur in the contrasting landforms. GPS location data and a machine learning technique were used to identify the seasonal pattern and the factors driving grazing pressure on a fenced ranch. Over both low-land and sand-dune landforms, the proportion of time that livestock spent on foraging increased from 63% in July to 67% in August and 69% in September, and non-foraging behavior decreased correspondingly. In low-land, the log-transformed average foraging density significantly increased from 0.61 (i.e., total foraging behaviors in 5 days measured at 50-s intervals per 10 × 10 m grid) in July to 0.66 in August and 0.88 in September, whereas there was no significant change on sand-dunes. From July to September, the relative area of low-land foraged by cattle accounted for 31%, 35%, and 36%, respectively, and in sand-dunes the proportions increased from 45% to 47% to 51%. In low-land, the foraging density was negatively correlated with biomass (P = .07), total digestible nutrients (P < .05), and crude protein (P = .06) and positively correlated with acid detergent fiber (P < .05), whereas no such relationships were observed in sand-dunes. Our results indicate that topographic features should be considered when managing livestock, especially during periods with adverse conditions of herbage quality and quantity.