Article: Cleptoparasitism and detritivory in dung beetle fossil brood balls from Patagonia, Argentina

Traces within traces is a new ichnological field that is meant to shed light on significative palaeoecological aspects. Dung beetle fossil brood balls (Coprinisphaera ispp.), from the Middle Eocene – Lower Miocene Sarmiento Formation of Patagonia, Argentina, show two different trace fossils excavated in its infillings and/or wall that reveal the presence and relationships among different components of past dung communities. Tombownichnus pepei n. isp. is represented by elongated pits, circular to elliptical in cross-section, occurring in the centre or beside ovoid mounds in the internal surface of the Coprinisphaera wall. These traces record the activity of cleptoparasites, such as other dung beetles or flies, whose larvae were probably carried passively with the dung for provisions. Tombownichnus pepei would represent the pupation chambers excavated by full grown larvae in the Coprinisphaera wall after completing their development inside provisioned dung. The other trace fossil, Lazaichnus fistulosus is represented by circular to subcircular borings occurring in Coprinisphaera walls, in connection with an internal gallery in their infillings. Its connection also with meniscate burrows and chambers in the surrounding palaeosol attributable to aestivation chambers of earthworms revealed that these organisms would have been active cleptoparasites or detritivores in dung beetle fossil brood balls.