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Maintaining multiple ecological functions ("multifunctionality") is crucial to sustain viable ecosystems. To date most studies on biodiversity-ecosystem functioning (BEF) have focused on single or few ecological functions and services. However, there is a critical need to evaluate how species and species assemblages affect multiple processes at the same time, and how these functions are interconnected. Dung beetles represent excellent model organisms because they are key contributors to several ecosystem functions. Using a novel method based on the application of N-enriched dung in a mesocosm field experiment, we assessed the role of dung beetles in regulating multiple aspects of nutrient cycling in alpine pastures over appropriate spatial (up to a soil depth of 20 cm) and temporal (up to 1 yr after dung application) scales. N isotope tracing allowed the evaluation of multiple interrelated ecosystem functions responsible for the cycling of dung-derived nitrogen (DDN) in the soil and vegetation. We also resolved the role of functional group identity and the importance of interactions among co-occurring species for sustaining multiple functions by focusing on two different dung beetle nesting strategies (tunnelers and dwellers). Species interactions were studied by contrasting mixed-species to single-species assemblages, and asking whether the former performed multiple functions better than the latter. Dung beetles influenced at least seven ecological functions by facilitating dung removal, transport of DDN into the soil, microbial ammonification and nitrification processes, uptake of DDN by plants, herbage growth, and changes in botanical composition. Tunnelers and dwellers were found to be similarly efficient for most functions, with differences based on the spatial and temporal scales over which the functions operated. Although mixed-species assemblages seemed to perform better than single-species, this outcome may be dependent on the context. Most importantly though, the different functions were found to be interconnected sequentially as reveled by analyzing N content in dung, soil and vegetation. Taken together, our current findings offer strong support for the contention that the link between biodiversity and ecosystem functions should be examined not function by function, but in terms of understanding multiple functions and how they interact with each other.

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