Ecological consequences of shrub encroachment in the grasslands of northern China

Context Shrub encroachment is a critical environmental issue in arid and semi-arid landscapes. However, due to the complicated effects of shrub characteristics and environmental variables, ecological consequences of shrub encroachment significantly differ in different regions. As a result, although the phenomenon of shrub encroachment is widespread in the grasslands of northern China, its ecological consequences and associated drivers have not been well documented at the regional scale. Objectives We explored the effects of shrub encroachment on ecosystem structures (herb richness and abundance) and functioning (above- and belowground biomass, and soil carbon and nitrogen content) and evaluated the relative importance of climatic variables (temperature and precipitation) and shrub characteristics (cover, area, and height) as drivers of these effects. Methods Based on field data from 53 sites that stretch across approximately 3000 km in northern China, we conducted the relative interaction intensity (RII) of the response variables between shrub patches and grassy matrix, and then used the structural equation modeling (SEM) to assess the relative importance of chosen drivers on the ecological consequences. Results Shrub encroachment increased the landscape heterogeneity by decreasing the richness, abundance and aboveground biomass of herbs, while by increasing the soil carbon and nitrogen content. The SEM explained 25–43% of the variation in shrub characteristics and 5–34% of the variation in the effects of shrub encroachment. Shrub height had greater effects on the RII of herb richness, soil carbon and soil nitrogen content, and the standardized total effects were 0.48, 0.25 and 0.18, respectively. Precipitation and temperature positively affected the impacts of shrub encroachment on herb richness, abundance, biomass, and multi-indexes. However, the direct effects of climate could be partially mediated by indirect effects via shrub characteristics. Conclusions The height, size and cover of shrub patches significantly influenced the community structures and functioning of grassland ecosystems. The increasing aridity could potentially exacerbate the negative consequences of shrub encroachment in arid regions. Our results also demonstrated that the effects of climate variables on grassland ecosystems could be mediated by shrub management.