Increased cover of native and exotic plants on the rims of harvester ant (Hymenoptera: Formicidae) nests under grazing and drought

Harvester ants create habitats along nest rims, which some plants use as refugia. These refugia can enhance ecosystem stability to disturbances like drought and grazing, but their potential role in invasion ecology is not yet tested. Here we examine the effects of drought and grazing on nest-rim refugia of 2 harvester ant species: Pogonomyrmex occidentals and P. rugosus. We selected 4 rangeland sites with high harvester ant nest densities in northern Arizona, USA, with pre-existing grazing exclosures adjacent to heavily grazed habitat. Our objective was to determine whether nest refugia were used by native or exotic plant species for each site and scenario of drought and grazing. We measured vegetation cover on nest surfaces, on nest rims, and at 3 distances (3, 5, and 10 m) from nests. At each site, we sampled 2 treatments (grazed/excluded) during 2 seasons (drought/monsoon). We found that nest rims increased vegetation cover compared with background levels at all sites and in almost all scenarios of treatment and season, indicating that nest rims provide important refugia for plants from drought and cattle grazing. In some cases, plants enhanced on nest rims were native grasses such as blue gramma (Bouteloua gracilis) or forbs such as sunflowers (Helianthus petiolaris). However, nest rims at all sites enhanced exotic species, particularly Russian thistle (Salsola tragus), purslane (Portulaca oleracea), and bull thistle (Cirsium vulgare). These results suggest that harvester ants play important roles in invasion ecology and restoration. We discuss potential mechanisms for why certain plant species use nest-rim refugia and how harvester ant nests contribute to plant community dynamics.