Palaeoecological aspects of some invertebrate trace fossils from the mid- to Upper Permian Middleton Formation (Adelaide Subgroup, Beaufort Group, Karoo Supergroup), Eastern Cape, South Africa

Ichnological and sedimentological analyses in the Eastern Cape allowed the first description of a Cochlichnus-dominated ichnofossil site from the mid- to Upper Permian Middleton Formation (Karoo Supergroup) in South Africa. The locality is within the uppermost Pristerognathus Assemblage Zone, a biostratigraphic interval characterized by a low vertebrate biodiversity at the turn of the mid- to Late Permian. Our field data indicates that the surficial bioturbation of very fine to fine-grained sand layers resulted from life activities of shallow infaunal and epifaunal invertebrates (possibly annelids, aquatic oligochaetes, nematodes, insect larvae) and fish. The morphology of the trails, their relationship to the substrate and the behaviour inferred from them indicate that the tracemakers developed a strategy that facilitated the optimization of low food resources in a permanently submerged freshwater setting. Combined ichnological and sedimentological evidence suggests a low-energy, freshwater lacustrine depositional environment, where occasional higher energy currents brought nutrients. Data also imply that colonization of these erratic event beds by opportunistic sediment-feeders was short-lived and followed by longer intervals of lower energy deposition under possibly poorly oxygenated conditions. We propose that these event beds as well as the sporadic red mudstones of the Middleton Formation may have formed during short-term, higher storm-frequency and dryer periods, signalling changes in the otherwise humid climate in this part of the main Karoo Basin during the mid- to Late Permian.

► Simple, low diversity, low abundance, shallow infaunal and epifaunal grazing traces. ► Ichnofossils represent an impoverished, permanently subaqueous Mermia ichnofacies. ► Associated with lacustrine event beds at the turn of the Middle to Late Permian. ► Middleton Formation lakes were deep enough for oxygen-stratification to occur.