The Paleozoic plant terrestrialization(PPT)represents a pivotal point in the evolutionary history of life on Earth. This process not only reshaped surficial geomorphological landscapes but also exerted on others biotic evolution throughout the Phanerozoic. Until now,how benthic animals respond to PPT across different depositional environments,as well as their roles in the construction of benthic ecosystems,have remained insufficiently understood. Based on the compilation and synthesis of a global trace fossil dataset,this study establishes a time-depositional environment analytical framework. Within this framework,we systematically compare the evolutionary patterns of ichnodiversity,ichnodisparity,ichnotiering,and benthic ecosystem engineering across transitional coastal,fluvial,and lacustrine settings during the process of PPT. Furthermore,we quantitatively assess the behavioral responses of benthic animals to PPT. The results show that from the Cambrian to the Permian,the behavioral evolution of benthic animals exhibits pronounced time-limited and heterogeneity. During the Ordovician,benthic animals in transitional coastal environments had already begun to show initial responses to PPT,whereas those in fluvial and lacustrine environments display a clear temporal lag. This discrepancy is closely associated with the development of plant root systems,the progression of PPT,and subsequent forest expansion. Overall,with the progressive improvement of substrate stability and organic matter availability in both continental and transitional settings driven by terrestrial plants,benthic animals gradually expanded their ecospace utilization,accompanied by increased behavioral complexity and enhanced ecosystem engineering. In turn,benthic animals exerted a positive feedback on PPT through sediment reworking and the modulation of matter and energy fluxes. In addition to the influence of terrestrial vegetation,environmental factors such as climate change and tectonic activity,and sea-level fluctuations also played significant roles in shaping benthic evolutionary trajectories. This study provides a novel ichnological perspective for understanding Paleozoic ecosystem evolution and biotic-sediment interactions.