Zhichun Lan

Evidence that acidification‐induced declines in plant diversity and productivity are mediated by changes in below‐ground communities and soil properties in a semi‐arid steppe

Summary 1. Anthropogenic acid deposition–induced soil acidification is one of the major threats to biodiversity, ecosystem functioning and services. Few studies, however, have explored in detail how above-ground changes in plant species richness and productivity resulting from soil acidification are mediated by effects on below-ground biota and soil properties. 2. To increase our understanding of this linkage, we collected data on below- and above-ground communities and soil properties in a 3-year field experiment with seven levels of acid addition rate to build-up broad intensities of soil acidification in the semi-arid Inner Mongolian grassland. 3. Acid addition directly elevated concentrations of soil Al 3+ ions, decreased the base cations Ca 2+ , Mg 2+ and Na + , and increased soil moisture and available phosphorus. Acid addition also appears to have altered the soil microbial community via changes in H + and Al 3+ ions and altered the nematode community via changes in H + ions and soil moisture. 4. The observed changes in soil N availability (i.e. net N mineralization, NO � -N and NH þ -N) could be explained by mediating changes in the H + and Al 3+ ions, microbial community (i.e. community structure, bacteria and fungi/bacteria as indicated by phospholipid fatty acids analysis) and the nematode community (i.e. total abundance, taxa richness and maturity index). 5. Declines in plant species richness and productivity were greater at high intensities of soil acidification in the second sampling year than in the first sampling year. The changes in plant community observed were mostly explained by soil nutrient pathways (e.g. N availability or base mineral cations), which were in turn regulated by the soil microbial or nematode communities as well as by the direct effects of the increase in H + or Al 3+ ions. 6. Synthesis. Our results suggest that the below-ground microbial and nematode communities are more sensitive to soil acidification than the plant communities are, and further that soil acidification–induced changes in plants are mediated by changes in below-ground communities and soil nutrients. These findings improve our understanding of the links between below- and above-ground communities in the Inner Mongolia grassland, especially in the context of anthropogenic acid enrichment.

Differential responses of plant functional trait to grazing between two contrasting dominant C3 and C4 species in a typical steppe of Inner Mongolia, China

Plant functional traits have been widely used to study the linkage between environmental drivers, trade-offs among different functions within a plant, and ecosystem structure and functioning. Here, the whole-plant traits, leaf morphological and physiological traits of two dominant species, Leymus chinensis (C3 perennial rhizome grass) and Cleistogenes squarrosa (C4 perennial bunchgrass), were studied in the Inner Mongolia grassland of China, with a grazing experiment including five stocking rates (0, 3.0, 4.5, 7.5, and 9.0 sheep/ha) in 2008 (wet year) and 2009 (dry year). Our results demonstrated that, for both species, the effects of stocking rate, year, and stocking rate × year on whole-plant traits and leaf morphological and physiological traits were highly significant in most cases. The differential responses of plant trait to variation in precipitation were caused by trait trade-offs between the wet and dry years. L. chinensis adopted the high N content and net photosynthetic rate (Pn) in the wet year but both the low N content and Pn in the dry year under grazed conditions. The trait trade-offs of C. squarrosa were characterized by high specific leaf area (SLA) and Pn in the dry year vs. low SLA and Pn in the wet year. Our findings also indicate that C. squarrosa is more resistant to grazing than L. chinensis in terms of avoidance and tolerance traits, particularly under heavy grazing pressure and in the dry year.

Testing mechanisms of N-enrichment- induced species loss in a semiarid Inner Mongolia grassland: critical thresholds and implications for long-term ecosystem responses

The increase in nutrient availability as a consequence of elevated nitrogen (N) deposition is an important component of global environmental change. This is likely to substantially affect the functioning and provisioning of ecosystem services by drylands, where water and N are often limited. We tested mechanisms of chronic N-enrichment-induced plant species loss in a 10-year field experiment with six levels of N addition rate. Our findings on a semi-arid grassland in Inner Mongolia demonstrated that: (i) species richness (SR) declined by 16 per cent even at low levels of additional N (1.75 g N m–2 yr−1), and 50–70% species were excluded from plots which received high N input (10.5–28 g N m−2 yr−1); (ii) the responses of SR and above-ground biomass (AGB) to N were greater in wet years than dry years; (iii) N addition increased the inter-annual variations in AGB, reduced the drought resistance of production and hence diminished ecosystem stability; (iv) the critical threshold for chronic N-enrichment-induced reduction in SR differed between common and rare species, and increased over the time of the experiment owing to the loss of the more sensitive species. These results clearly indicate that both abundance and functional trait-based mechanisms operate simultaneously on N-induced species loss. The low initial abundance and low above-ground competitive ability may be attributable to the loss of rare species. However, shift from below-ground competition to above-ground competition and recruitment limitation are likely to be the key mechanisms for the loss of abundant species, with soil acidification being less important. Our results have important implications for understanding the impacts of N deposition and global climatic change (e.g. change in precipitation regimes) on biodiversity and ecosystem services of the Inner Mongolian grassland and beyond.

Soil acidification exerts a greater control on soil respiration than soil nitrogen availability in grasslands subjected to long‐term nitrogen enrichment

Summary Terrestrial ecosystems worldwide are receiving increasing amounts of biologically reactive nitrogen (N) as a consequence of anthropogenic activities. This intended or unintended fertilization can have a wide-range of impacts on biotic communities and hence on soil respiration. Reduction in below-ground carbon (C) allocation induced by high N availability has been assumed to be a major mechanism determining the effects of N enrichment on soil respiration. In addition to increasing available N, however, N enrichment causes soil acidification, which may also affect root and microbial activities. The relative importance of increased N availability vs. soil acidification on soil respiration in natural ecosystems experiencing N enrichment is unclear. We conducted a 12-year N enrichment experiment and a 4-year complementary acid addition experiment in a semi-arid Inner Mongolian grassland. We found that N enrichment had contrasting effects on root and microbial respiration. N enrichment significantly increased root biomass, root N content and specific root respiration, thereby promoting root respiration. In contrast, N enrichment significantly suppressed microbial respiration likely by reducing total microbial biomass and changing the microbial community composition. The effect on root activities was due to both soil acidity and increased available N, while the effect on microbes primarily stemmed from soil acidity, which was further confirmed by results from the acid addition experiment. Our results indicate that soil acidification exerts a greater control than soil N availability on soil respiration in grasslands experiencing long-term N enrichment. These findings suggest that N-induced soil acidification should be included in predicting terrestrial ecosystem C balance under future N deposition scenarios.

Scale-Dependent Effects of Grazing on Plant C: N: P Stoichiometry and Linkages to Ecosystem Functioning in the Inner Mongolia Grassland

Background Livestock grazing is the most prevalent land use of grasslands worldwide. The effects of grazing on plant C, N, P contents and stoichiometry across hierarchical levels, however, have rarely been studied; particularly whether the effects are mediated by resource availability and the underpinning mechanisms remain largely unclear. Methodology/Principal Findings Using a multi-organization-level approach, we examined the effects of grazing on the C, N, and P contents and stoichiometry in plant tissues (leaves and roots) and linkages to ecosystem functioning across three vegetation types (meadow, meadow steppe, and typical steppe) in the Inner Mongolia grassland, China. Our results showed that the effects of grazing on the C, N, and P contents and stoichiometry in leaves and roots differed substantially among vegetation types and across different hierarchical levels (species, functional group, and vegetation type levels). The magnitude of positive effects of grazing on leaf N and P contents increased progressively along the hierarchy of organizational levels in the meadow, whereas its negative effect on leaf N content decreased considerably along hierarchical levels in both the typical and meadow steppes. Grazing increased N and P allocation to aboveground in the meadow, while greater N and P allocation to belowground was found in the typical and meadow steppes. The differences in soil properties, plant trait-based resource use strategies, tolerance or defense strategies to grazing, and shifts in functional group composition are likely to be the key mechanisms for the observed patterns among vegetation types. Conclusions/Significance Our findings suggest that the enhanced vegetation-type-level N contents by grazing and species compensatory feedbacks may be insufficient to prevent widespread declines in primary productivity in the Inner Mongolia grassland. Hence, it is essential to reduce the currently high stocking rates and restore the vast degraded steppes for sustainable development of arid and semiarid grasslands.

Functional trait responses to grazing are mediated by soil moisture and plant functional group identity

Abundant evidence has shown that grazing alters plant functional traits, community structure and ecosystem functioning of grasslands. Few studies, however, have tested how plant responses to grazing are mediated by resource availability and plant functional group identity. We examined the effects of grazing on functional traits across a broad range of species along a soil moisture gradient in Inner Mongolia grassland. Our results showed that trait syndromes of plant size (individual biomass) and shoot growth (leaf N content and leaf density) distinguished plant species responses to grazing. The effects of grazing on functional traits were mediated by soil moisture and dependent on functional group identity. For most species, grazing decreased plant height but increased leaf N and specific leaf area (SLA) along the moisture gradient. Grazing enhanced the community-weighted attributes (leaf NCWM and SLACWM), which were triggered mainly by the positive trait responses of annuals and biennials and perennial grasses, and increased relative abundance of perennial forbs. Our results suggest that grazing-induced species turnover and increased intraspecific trait variability are two drivers for the observed changes in community weighted attributes. The dominant perennial bunchgrasses exhibited mixed tolerance–resistance strategies to grazing and mixed acquisitive–conservative strategies in resource utilization.

Scale-dependent patterns and mechanisms of grazing-induced biodiversity loss: evidence from a field manipulation experiment in semiarid steppe

ContextAlthough many studies have demonstrated that grazing may increase or decrease plant diversity of grasslands at small scales, few studies have examined the patterns and mechanisms of grazing effects on biodiversity across multiple scales.ObjectiveOur study tested the scale dependence of grazing effects on plant diversity based a 7-year grazing manipulation experiment with seven levels of grazing intensity (0–9 sheep ha−1) in a typical steppe of the Inner Mongolia grassland.MethodsSpecies area relationships (SARs) were used to analyze the scale dependence of species loss. SAR decomposition approaches were followed to examine the contribution of four potential mechanisms to changes in the slope of SARs, including species aggregation, overall species richness, total number of individuals, and species abundance distribution.ResultsThe proportional species loss increased with sampling area (4–1024 m2), which was evidenced by decreasing intercepts and slopes of SARs with grazing. Reduction in the slope of SARs was mainly caused by changes in overall species richness and species abundance distribution, with the relative minor effect of changes in number of individuals.ConclusionsThe negative effect of grazing on overall species richness was mainly attributed to the loss of grazing-sensitive rare species from species pool. Compared with flat systems, plant diversity in slope systems was more sensitive to grazing at low intensities. However, the responses of plant diversity to grazing tended to converge between the flat and slope systems at high levels of grazing intensity. Our study has important implications for adaptive ecosystem management and biodiversity conservation in arid and semiarid grasslands.

Testing the scaling effects and mechanisms of N‐induced biodiversity loss: evidence from a decade‐long grassland experiment

Summary 1. Although extensive studies demonstrate that nitrogen (N) enrichment frequently reduces plant diversity within small quadrats (0.5– 4m 2 ), only a few studies have evaluated N effects on biodiversity across different spatial scales. 2. We conducted the first experimental test of the scale dependence of N effects on species richness from a 10-year N treatment (1.75–28 g N m � 2 year � 1 ) in a typical steppe. We used species–area relationship (SAR) to analyse the scale dependence of species loss with power model S = cA z (S is species number, A is area, c is intercept, and z is slope). 3. Absolute species loss decreased at sampling area > 8 m 2 . Proportional species loss (compared to control) decreased and critical threshold (Ncrit) for biodiversity losses increased with sampling areas. 4. These scale dependences were quantified as increasing slope (z-value) of SAR with N addition. Through SAR decomposition, we found that this overall positive effect was in response to positive effects of changes to the species abundance distribution over negative effects of overall species richness losses. 5. Synthesis. As nitrogen (N) enrichment typically occurs at scales much larger than individual plots, understanding how N enrichment affects the scaling patterns of biodiversity is necessary for biodiversity conservation and ecosystem management in response to anthropogenic N deposition.