Ichnology of a Middle Devonian regression: Environment, terrestrialization and true substrate controls on trace fossils of the Hangman Sandstone Formation, SW England

The Eifelian-aged Hangman Sandstone Formation of southwest England is a unit of the ‘Old Red Sandstone’ that has traditionally been considered relatively barren of trace fossils. Following recent investigations, we here show it to contain the most diverse Middle Devonian non-marine ichnofauna known globally. The fluvial-lacustrine facies of its constituent Trentishoe Member contain representatives of 21 ichnogenera including Archaeonassa, Arenicolites, Beaconites, Bifungites, Circulichnis, Cruziana, Diplichnites, ?Gluckstadella, Gordia, ?Halimededes, ?Lockeia, Merostomichnites, Palmichnium, Petalichnus, Planolites, Polarichnus, Rusophycus, Siskemia, Spirophyton, Steinsfjordichnus, and Taenidium, as well as several microbially induced sedimentary structures. The transitional marine-influenced facies of the underlying Hollowbrook Member are less extensively exposed but contain four additional unique ichnogenera in the form of Cochlichnus, Halopoa, Phycodes, and Teichichnus, as well as further examples of Cruziana, Planolites and Taenidium. A revised ichnological assessment of the unit reveals several notable features, including the youngest known example of the arthropod refugia trace, Polarichnus, as well as a transitional trace fossil form of Diplichnites-Beaconites, demonstrating that the latter was produced by arthropods. As the Hangman Sandstone Formation records the culmination of a marine regression, its ichnofauna can be compared with that of the conformably underlying Lynton Formation, which was deposited in a sandy marine shelf setting. The ichnological differences between these units are acute, with 88% of the Hangman Sandstone Formation trace fossils being exclusive to non-marine or transitional facies. This observation sheds light on the terrestrialization process and indicates that the segregation of marine and non-marine trace fossil communities had accelerated by the Middle Devonian. Internal variability in ichnological signatures within the Hangman Sandstone Formation is also pronounced, with greater ichnodiversity and ichnodisparity in distal fluvio-lacustrine facies than in proximal fluvial facies. By viewing the depositional environment of the unit as a regionally extensive distributive fluvial system, this variability can be partly explained by a bias arising from the distribution of true substrates (bedding planes that demonstrably preserve an ancient sediment-air interface) in the unit, with such phenomena being more readily preserved in the lower energy outer reaches of the depositional system.