Complex behavioural pattern as an aid to identify the producer of Zoophycos from the Middle Permian of Oman

The trace fossil Zoophycos is abundant in transgressive, shallow marine carbonates in the Middle Permian (Wordian) Khuff Formation of the Huqf-Haushi Uplift of Interior Oman. It often occurs as part of a complex (compound) trace fossil that comprises two integrated elements: (i) irregular galleries with straight to gently curved tunnels and interconnected shafts, and (ii) simple planar to complex spreiten structures with a marginal tube (Zoophycos). The galleries are characterized by irregularly winding, dichotomous branching, large variation in shape and size and circular to elliptical vertical cross-sections. Zoophycos consists of spreiten with a marginal tube, either originating as a simple lobe from the convex segment of a curved tunnel, or forming more complex, subcircular, spreiten systems parallel to bedding. The spreiten were formed by simple strip mining, where the animal defecated without producing faecal pellets. U-shaped marginal tubes indicate that the burrows were well aerated. The complex trace fossil points to combined dwelling and deposit-feeding behaviour, with irregular galleries in the firm substrate and Zoophycos spreiten in the softground below it. It can be assumed that the animal used the open tunnel system mainly for dwelling (domichnion) and possibly suspension feeding, but occasionally changed to deposit feeding while creating the spreiten (fodinichnion). The integration of the irregular galleries (tunnels and interconnected shafts) with the marginal tubes of Zoophycos suggests the same producer for this compound trace fossil. Many modern polychaetes produce very similar galleries within firm and soft substrates, and polychaetes are therefore interpreted as the most likely producers. Similarities between Permian and Triassic Zoophycos suggest comparable trace making behaviour before and after the end-Permian mass extinction.