The role of dung beetles as a secondary seed disperser after dispersal by frugivore mammals in a temperate deciduous forest

We studied the effects of dung beetles on the fates of endozoochorous seeds of five species (Prunus jamasakura, Prunus verecunda, Prunus grayana, Swida controversa, and Vitis coignetiae) in a temperate deciduous forest in Japan during 2004–2006. In field experiments using dung of the Asiatic black bear (Ursus thibetanus), we investigated the depths that dung beetles (Onthophagus atripennis, Onthophagus lenzii, and Phelotrupes auratus) buried seeds (4.8–6.8 mm diameter) and plastic markers (2 or 5 mm diameter), the levels of predation on buried and unburied seeds, and germination rates of seeds buried to different depths. All three species buried the 2-mm markers, but only P. auratus buried the seeds and 5-mm markers. There were seasonal differences in mean seed burial rates (range, 27–51%) and depths (range, 1–27 mm). Significantly more seeds were buried in June, July, and September than in August or October, and the mean burial depth was significantly deeper in June and July. Most seeds and markers were buried to a 3–6 cm depth. Germination of seeds that were positioned at depths of 1–4 cm was significantly greater than that of seeds left on the surface or buried at greater depths. Buried seeds were less likely to disappear than seeds at the surface, which may reflect differential predation. These results suggested that dung beetles, especially P. auratus, acted as a secondary seed disperser that affected the survival and distribution of seeds dispersed by a frugivore.