Fig. 1: Geographic location of sampling sites across 4000 km of grasslands.

a Inset in the bottom right depicts the spread of mean annual temperature (MAT) and mean annual precipitation (MAP) across these sites. b Schematic diagram of shrub patches spaced with grassland patches in shrub-invaded grasslands. The blue dotted area denotes a shrub patch, and the orange dotted area illustrates an interspaced grassland patch.