About 200 Zoophycos specimens, including 90 specimens studied in detail, have been analysed in the continuous Upper Cretaceous–Lower Miocene pelagic sedimentary type sections of the Gubbio area (the Contessa Highway, Contessa Quarry and Bottaccione sections, Northern Apennines). The sediments are reddish to grey limestones and marls of the Scaglia Group and marls with volcaniclastic deposits of the Bisciaro Formation. The aim was to examine the evolutionary trend of what is probably the most debated trace fossil of all time, from the Upper Cretaceous to Lower Miocene. Despite having been found in beds ranging from the Cambrian to the present, no consensus has been reached regarding mode of construction, tracemaker or ethological explanation for Zoophycos. Four Zoophycos morphotypes are recognized at Gubbio showing variations of major and minor lamellae, apex, lobes and whorls: the Cretaceous–Eocene cone-shaped type 1, the Upper Eocene–Middle Oligocene helicoidal type 2, the Oligocene lobate type 3 and the Upper Oligocene–Lower Miocene flat type 4. The very high ichnodensity in some beds (hundreds of specimens in discrete levels of the Bisciaro Formation, now destroyed by quarrying) seems to find explanation in abnormal concentrations of phytodetritus and organic matter on the seafloor in some periods. This very high abundance in discrete levels reflects a change in sedimentation and seafloor conditions at pre-flysch deposition. Due to such high ichnodensity, many adjacent specimens display deformed outer margins. Taphonomic analysis shows a variation of whorls, laminae and U-shaped lobes, reflecting ontoge- netic development of the tracemaker(s) (?sipunculid worms).
Monaco, P., Bracchini, L., Rodr ıguez-Tovar, F.J., Uchman, A. & Coccioni, R. 2016: Evolutionary trend of Zoophycos morphotypes from Upper Cretaceous–Lower Miocene in the type pelagic sections of Gubbio, central Italy. Lethaia, DOI: 10.1111/ let.12175.