Temporal coexistence of dung-dweller and soil-digger dung beetles (Coleoptera, Scarabaeoidea) in contrasting Mediterranean habitats

The western part of the Mediterranean basin is a transitional biogeographical region for the distribution of the representatives of the main guilds of dung beetles; towards the south, Aphodiinae (dung-dwellers) become scarce, whereas northwards Scarabaeinae (soil-diggers) progressively disappear. The number of species in local dung beetle assemblages is enhanced by this double faunistic contribution. Annual dung beetle assemblages were sampled in two sub-Mediterranean sites, which differed by 600 m in elevation, in order to determine the phenological dynamics related to the way of using dung (dung-dwellers/Aphodiinae vs. soil-diggers/Scarabaeinae and Geotrupinae). Aphodiids were active all year round, although they were affected by summer drought and, at high elevation, by the length of the cold season. This reduced activity was related to an impoverishment of Aphodiinae and to reduced temporal segregation between species. In contrast, soil-diggers were not active all year round and showed different species assemblages in the two sites. An extension of the activity period of these beetles was observed due to the occurrence of cold resistant species at high elevation. Our results suggested that the occurrence of soil-diggers seemingly did not affect the seasonality of dung-dwellers; their local abundance showed no negative correlation and, most importantly, phenological differences between dung-dwellers were always significantly higher than the seasonal differences between dwellers and diggers.