Aggregated distribution of resources creates competition refuges for rainforest dung beetles

This study examines the effects of resource distribution on colonisation, intra- and interspecific aggregation, and the occurrence of low-density, competition refuges for tropical dung beetles. In field experiments from central Peru, using dung pats (resource patches) of different volumes, the numbers of interacting species and total beetle biomass at individual pats increased with increasing pat volume. In two of three separate experiments (including an experiment that also varied patch density), this represented a decrease in the biomass of beetles per unit volume (biomass-density) at larger patches. The numbers of interacting tunneller species and tunneller biomass-density were also related to the distance between pats (patch density) in one of two experiments with constant numbers of pats. Closely positioned pats had generally fewer interacting species and a lower biomass-density of beetles. For the most abundant Dichotomius species, interspecific associations increased as distances increased between dung pats. The numbers of interacting species and biomass-density declined linearly under the combined effects of increasing patch density and local patch abundance in 25 m² plots. In experimentally placed grids with large numbers of pats, colonisation of pats at the edge of the grids was generally higher than at the centre of the grids for tunnellers and Eurysternus spp. but not for ball rollers; however, at least tunnellers did not readjust to avoid patches with high densities of competitors. These results indicate that an aggregated distribution of dung and natural variability in patch size contribute to species coexistence by creating low-density refuges for weaker competitors.