Trait-mediated responses and co-occurrence patterns of breeding steppe birds under human footprint in Inner Mongolia

Grasslands are among the world's most threatened ecosystems, and steppe birds face increasing risks from human activities. This study investigates how human impacts affect the distribution and community structure of breeding steppe birds in Inner Mongolia, a biodiversity hotspot in Asia. We conducted standardized point-count surveys across a gradient from intact grasslands to urbanized areas, integrating species occurrence data, functional traits and the Human Footprint Index (HFI). Using Generalized Linear Models (GLMs) and Conditional Random Fields (CRFs), we assessed trait-environment interactions and shifts in species associations. Our results indicate that the HFI significantly affects bird communities. Habitat specialists, such as Mongolian Lark (Melanocorypha mongolica), showed negative responses, whereas generalists like Eurasian Tree Sparrow (Passer montanus) thrived in disturbed areas. Trait-based analysis showed that species with larger body mass and specialized diets were negatively associated with HFI, whereas those linked to human-modified habitats exhibiting strong positive associations. In areas with high human footprint, co-occurrence networks grew more polarized: specialists faced intensified competition, while species with positive HFI responses formed stronger positive associations. CRF models indicated that human activities restructure species interactions, favoring generalists and simplifying community dynamics. These findings highlight the dual role of human impact in supporting some species while threatening specialists, potentially driving biotic homogenization. Our study emphasizes the need for conservation strategies that protect vulnerable species and manage those that thrive in human-altered environments. By linking traits and interactions to human impacts, this study provides a framework for identifying at-risk species and guiding conservation in the Anthropocene.