What do dung beetles eat?

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Co‐declining mammals and dung beetles: an impending ecological cascade (2009, background): “Drawing firm conclusions regarding the impacts of reduced dung diversity in persistently hunted forests is complicated by the dearth of empirical information on the feeding ecology of individual dung beetle species (Holter and Scholtz 2007).”
Herbivore dung as food for dung beetles: elementary coprology for entomologists (2016, background): “What is proposed here, as well as by Holter and Scholtz (2007), is that dung beetles subsist on the dead and alive microbial biomass in fresh dung, possibly supplemented by small amounts of other particulate debris from the herbivore’s gut.”
A brown‐world cascade in the dung decomposer food web of an alpine meadow: effects of predator interactions and warming (2011, background): “Adults of coprophagous beetles have mouth- parts that allow them to feed only on dung liquids and smaller dung particles, and meanwhile filter out large and indigestible plant fragments (Holter and Scholtz 2007).”