Stars of the aftermath: Asteriacites beds from the Lower Triassic of the Carnic Alps (Werfen Formation, Sauris di Sopra), Italy

Trace fossils are important evidence of benthic activity, but they have received less study than body fossils for investigating the aftermath of the end-Permian extinction. There is therefore a need to document Lower Triassic ichnofaunas to understand their significance with respect to the end-Permian crisis. In light of this need, this paper describes a novel Lower Triassic ichnosite at Mount Pallone (Carnic Alps, Italy), where the Campil Member (Smithian) of the Werfen Formation (Griesbachian-Spathian) presents an abundant ichnofauna characterized by excellent preservation and low diversity. Documented ichnogenera include Asteriacites lumbricalis, Gyrochorte comosa, Diplocraterion habichi and Planolites beverleyensis. The ichnofaunal composition and the bioturbation style suggest a marginal marine paleoenvironment ranging from intertidal to shallow subtidal settings. Storm influence, hydrodynamic energy, sedimentation rate, freshwater input and/or water temperature played an important role in structuring the benthic ecosystem. Dense (300 specimens/m2) aggregations of the trace fossil Asteriacites lumbricalis reveals social behavior of their inferred brittlestar producers (Echinodermata, Ophiuroidea). In line with modern brittlestar beds, social behavior provided significant advantage because raised arms of brittlestars dampened hydrodynamic energy. This study suggests that Asteriacites beds may be considered ichnological proxies for marine settings, low bioturbation intensity, shallow tiering, high sedimentation rate and/or event-bed deposition, significant levels of hydrodynamic energy, and low predation pressure. The studied ichnofauna reflects stressed environmental conditions, but it is unclear whether this reflects local brackish conditions ('Gazpacho model') or global hot temperatures ('Hot Soup model').