Biodiversity and soil pH regulate the recovery of ecosystem multifunctionality during secondary succession of abandoned croplands in northern China

The 'Grain-for-Green' program in China provides a valuable opportunity to investigate whether spontaneous restoration can reverse the deterioration of grassland ecosystem functions. Previous studies have focused on individual ecosystem functions, but the response of and mechanisms driving variation in ecosystem multifunctionality (EMF) during restoration are poorly understood. Here, we quantified EMF using productivity, nutrient cycling, and water regulation functions along abandoned croplands in a recovery chronosequence (5, 15 and 20 years) and in natural grasslands in the desert steppe and typical steppe. We also analyzed the effects of plant and microbial diversity and an abiotic factor (soil pH) on EMF. Our results showed that EMF increased gradually concomitant with recovery time, shifting toward EMF values comparable to those in natural grasslands in both desert and typical steppe. Similar results were found for the productivity function, the water regulation function, and soil organic carbon. However, even after 20 years of restoration, EMF did not reach the levels observed in natural grasslands. Structural equation modeling showed that the driving mechanisms of EMF differed between the two steppe types. Specifically, in the desert steppe, plant diversity, especially the diversity of perennial graminoids and perennial herbs, had a positive effect on EMF, but in the typical steppe, soil bacterial diversity had a negative effect, while soil pH had a positive effect on EMF. Our results demonstrated that spontaneous grassland restoration effectively enhanced EMF, and emphasized the importance of biodiversity and soil pH in regulating EMF during secondary succession. This work provides important insights for grassland ecosystem management in arid and semi-arid regions.