Intra- and interspecific aggregation of north temperate dung beetles on standardised and natural dung pads: The influence of spatial scale

1. The spatial distribution of north temperate dung beetles (Aphodius, Sphaeridium, and Margarinotus) was investigated at three spatial scales (scale a between all pads - large spatial scale, small patch size; scale b between block - large spatial scale, large patch size; scalec within block - small spatial scale, small patch size) in the field using standardised and naturally dropped dung pads that varied in size and volume. 2. Results indicated that all the major colonising species of dung beetle expressed strong intraspecific aggregation, while interspecific aggregation, though present, was of a lesser magnitude. 3. In both controlled and natural dung treatments, intra- and interspecific aggregation decreased with increasing patch size (from pad to block) and also decreased with decreasing spatial scale (from whole plot to within-block scale). 4. The data also suggested that intraspecific aggregation was more important than interspecific aggregation in the distribution of adult dung beetles in natural and standardised dung pads and hence has the potential to facilitate coexistence. 5. Intra- and interspecific aggregation was greater in the natural dung compared with that of the standardised dung, and the effect of pad size partly explains this phenomenon. 6. The within-patch spatial distribution of dung beetles observed in the natural and standardised dung was possibly mediated through within-patch differences between the two treatments. Six confounding factors could explain this difference and these factors are discussed in relation to resource utilisation by north temperate dung beetles.