Spatial heterogeneity increases diversity and stability in grassland bird communities.

Grasslands are inherently dynamic in space and time, evolving with frequent disturbance from fire and herbivores. As a consequence of human actions, many remaining grasslands have become homogenous, which has led to reduced ecosystem function, biodiversity loss, and decreased ecological services. Previous research has shown that restoring inherent heterogeneity to grasslands can increase avian diversity, but the amount of heterogeneity (i.e., number of patches or fire return interval) and the impact on avian community stability have yet to be investigated. We used a unique landscape-level design to examine avian response to interacting fire and grazing across multiple experimental landscapes that represented a gradient of fire- and grazing-dependent heterogeneity. We used seven landscapes (430-980 ha; x = 627 ha) with varying levels of patchiness ranging from annually burned (one single patch) with spring-only fires to a four-year fire return interval with spring and summer fires (eight patches). This design created a range of heterogeneity as a result of pyric herbivory, an ecological process in which fire and grazing are allowed to interact in space and time. We found that greater heterogeneity across experimental landscapes resulted in increased avian diversity and stability over time. An index of bird community change, quantified as the sum of the range of detrended correspondence analysis axis site scores, was nearly four times greater in the most homogenous experimental landscape when compared to the most heterogeneous experimental landscape. Species responses were consistently positively associated with increased heterogeneity at the landscape scale, and within-experimental-landscape responses were most often related to litter cover, litter accumulation, and vegetation height. We conclude that increased fire- and grazig-dependent heterogeneity can result in high variability in the bird community at finer, transect scales, but increased diversity and stability at broad landscape scales. We recommend that future management efforts in rangelands focus on restored disturbance processes to increase heterogeneity and improve grassland bird conservation.