Burrowing Richardson's ground squirrels affect plant seedling assemblages via environmental but not seed bank changes

In grassland ecosystems, burrowing mammals create disturbances, providing habitat for animal species and increasing plant community diversity. We investigated whether seedling assemblages on Richardson's ground squirrel Urocitellus richardsonii mounds result from seed rearrangement or environmental changes that favor germination of certain species over others. To test whether ground squirrels rearrange the seed bank by burrowing, we compared seed compositions among mounds, burrows, and undisturbed soil. To test whether ground squirrels change environmental conditions, we compared soil nitrate and bare ground cover on and off mounds. We also compared seedlings that germinated on mounds with seedlings that germinated on artificial disturbances from which we removed aboveground vegetation. Soil nitrate and bare ground cover were significantly higher on mounds than artificial disturbances. While seed richness and abundance did not differ among mounds, burrows, and undisturbed soil, seedling richness was reduced on mounds relative to artificial disturbances. Burrowing disturbance favors seedlings that can capitalize on bare ground availability but are less able to immobilize nitrate, as opposed to perennial species that immobilize more nitrate but take longer to establish. Our results suggest that Richardson's ground squirrels act as ecosystem engineers, although future research following succession on ground squirrel mounds is necessary to understand how they influence plant communities past the seedling stage.