Recent efforts to apply sustainability concepts to entire landscapes have seen increasing interest in approaches that connect socioeconomic and biophysical systems. Evaluating these connections through a cultural ecosystem services lens clarifies how different spatiotemporal scales and levels of organisation influence the production of cultural benefits. Currently, however, the effects of multi-level and multi-scale ecological variation on the production of cultural benefits have not yet been disentangled. Objectives To quantify the amount of variation in cultural ecosystem service provision by birds to birders that is due to landscape-level attributes. Methods We used data from 293 birding routes and 101 different birders in South African National Parks to explore the general relationships between birder responses to bird species and environmental conditions, bird-related observations, the biophysical attributes of the landscape and their effect on bird-related cultural benefits. Results Biophysical attributes (particularly biome, vegetation type, and variance in elevation) significantly increased the percentage of variance explained in birder benefits from 57 to 65%, demonstrating that birder benefits are derived from multi-level (birds to ecosystems) and multi-scale (site to landscape) social and ecological interactions. Conclusions Landscape attributes influence people's perceptions of cultural ecosystem service provision by individual species. Recognition of the complex, localised and inextricable linkage of cultural ecosystem services to biophysical attributes can improve our understanding of the landscape characteristics that affect the supply and demand of cultural ecosystem services
