Tropical and subtropical grasslands, savannas, and other dryland ecosystems are geographically extensive and provide critical support for human livelihoods (Parr et al., 2014). In these ecosystems, woody plant cover and abundance have been increasing in response to changes in climate, fire frequency, herbivory, and atmospheric CO 2 concentration during the past century (Archer et al., 2017). This geographically widespread phenomenon has dramatic impacts on multiple ecosystem functions and services (Eldridge et al., 2011). For example, numerous studies conducted across a wide range of ecosystems have shown that soil nutrient