Zoophycos is widely distributed in the marine strata of the Middle Permian Ruteh Formation in the Alborz Mountain, Iran. The investigation of the Zoophycos, along with environmental variables is a useful tool for interpretation of the palaeoenvironmental and sequence stratigraphy analysis. The petrographic observations led to the identification of ten facies in four facies belts including tidal flat, lagoon, shoal, and open marine, deposited on a homoclinal ramp. Moreover, two third-order depositional sequences were recognized in response to the sea-level fluctuations within the Ruteh Formation. Detailed studies of the sequence stratigraphy revealed a relationship between the occurrences of Zoophycos and changes in the hydrodynamic condition in the basin. It appears that Zoophycos has been influenced by the ecological and palaeoenvironmental parameters, such as sedimentation rate, nutrient supply, oxygen, wave base, and substrate in the shallow to deep environments. Based on the sedimentological and ichnological analysis, Zoophycos has been formed with various dimensions, morphology, fillings, and densities together with rising and falling in the sea-level. The trace-maker has followed an opportunistic strategy in the unstable conditions of shallow environments, whereas it has chosen a k-selected strategy in more stable deep environments. Additionally, variability in Zoophycos illustrates, how the trace-maker adopted itself with environmental sequences. This reason, owing to optimal conditions, has caused that the abundance of Zoophycos was high in the Transgressive System Tracts (TST). Evidence shows that the response of Zoophycos to the ecological properties of the environment usually has deposit-feeder and chemosymbiosis behaviours.

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