The soil macrofauna plays an important role in the carbon and nitrogen cycle of terrestrial ecosystems. In order to gain more insight into the role of the intestinal microbiota in transformation and mineralization of organic matter during gut passage, we characterized the physicochemical conditions, microbial activities, and community structure in the gut of our model organism, the humus-feeding larva of the cetoniid beetle Pachnoda ephippiata. Microsensor measurements revealed an extreme alkalinity in the midgut, with highest values (pH > 10) between the second and third crown of midgut ceca. Both midgut and hindgut were largely anoxic, but despite the high pH, the redox potential of the midgut content was surprisingly high even in the largest instar. However, reducing conditions prevailed in the hindgut paunch of all instars (E h ϳ ؊100 mV). Both gut compartments possessed a pronounced gut microbiota, with highest numbers in the hindgut, and microbial fermentation products were present in high concentrations The soil macrofauna plays an important role in the carbon and nitrogen cycle of terrestrial ecosystems (18,41,62). The intestinal tracts of litter-feeding and humivorous soil macroinvertebrates are favorable habitats for microorganisms and typically harbor a dense and active gut microbiota. The major function commonly attributed to the microorganisms in the guts of such animals is the depolymerization and fermentative breakdown of the cellulosic or lignocellulosic component of their diet, which leads to degradation products that can be resorbed by the host. This is supported by the high concentrations of microbial fermentation products and by the presence of fermentative bacteria and protozoa, accompanied by obligately anaerobic homoacetogenic and methanogenic microorganisms, in the guts of such animals. However, the extent and importance of such processes and their specific function in the nutrition of the host are scarcely understood (for reviews, see references 8, 9, 11, 12, 14, 36, and 44).In the case of soil-feeding termites, host factors such as the extreme alkalinity of the anterior hindgut and the influx of oxygen seem to play a key role in sequestering organic matter from the inorganic soil matrix (37). The decrease of molecular weight and increase in solubility resulting from alkaline extraction and chemical oxidation render the organic matter accessible for digestion in subsequent, less-alkaline compartments (34, 35). Alkaline gut regions are encountered also in many representatives of other insect orders and seem to be connected with the dietary preferences of the respective taxa (Coleoptera, Diptera, and Lepidoptera; see references 15 and 32). Comparing several beetle larvae feeding on a lignocellulosic diet, Grayson (30) already had pointed out an apparent correlation between the degree of humification of the diet and the alkalinity of the intestinal tract. The highest pH values among beetle larvae were encountered among the Scarabaeidae (50,52,58,60), which comprise species from humivorous, ...