Woody-plant encroachment: Precipitation, herbivory, and grass-competition interact to affect shrub recruitment.

Woody-plant encroachment is a global phenomenon that has been affecting the southwestern United States since the late 1800s. Drought, overgrazing, herbivory, and competition between grasses and shrub seedlings have been hypothesized as the main drivers of shrub establishment. However, there is limited knowledge about the interactions among these drivers. Using a rainfall manipulation system and various herbivore exclosures, we tested hypotheses about how precipitation (PPT), competition between grasses and shrub seedlings, and predation affect the germination and first-year survival of mesquite (Prosopis glandulosa), a shrub that has encroached in Southern Great Plains and Chihuahuan Desert grasslands. We found that mesquite germination and survival (1) increased with increasing PPT, then saturated at about the mean growing season PPT level, (2) that competition between grasses and shrub seedlings had no effect on either germination or survival, and (3) that herbivory by small mammals decreased seedling establishment and survival, while ant granivory showed no effect. In addition to its direct positive effect on survival, PPT had an indirect negative effect via increasing small mammal activity. Current models predict a decrease in PPT in the southwestern United States with increased frequency of extreme events. The non-linear nature of PPT effects on Mesquite recruitment suggests asymmetric responses, wherein drought has a relatively greater negative effect than the positive effect of wet years. Indirect effects of PPT, through its effects on small mammal abundance, highlight the importance of accounting for interactions between biotic and abiotic drivers of shrub encroachment. This study provides quantitative basis for developing tools that can inform effective shrub management strategies in grasslands and savannas.