Earliest evidence of granivory from China (Shanxi Formation) points to seeds as a food source and nursing habitat for insects in the earliest Permian humid tropical forests of Cathaysia

Three types of plant-insect interactions are identified on seeds from the lower Permian (Asselian) Shanxi and lower Shihhotse formations of the Taiyuan district, North China. This enhances the relatively meagre fossil record of seed predation in global late Paleozoic floras, adding the earliest record of granivory from Cathaysia. The dispersed seeds cannot be attributed with confidence to any particular plant group, but associated fossil leaves belong to a broad spectrum of plants, including Medullosales, Cycadales, Noeggerathiales, Gigantopteridales, Cordaitales, and Voltziales. Among 85 analysed seeds, six showed evidence of predation, referable to three damage types: DT074 and two new damage types that will be added to the forthcoming version of the fossil damage guide (DT274, DT430). These damage features indicate novel strategies of seed exploitation in the earliest Permian of China. The causal agents of the seed herbivory are difficult to resolve with certainty, but possible culprits include representatives of Palaeodictyopteroidea, although we cannot exclude other groups, such as Dictyoptera, Odonatoptera, Archaeorthoptera, Hemipteroidea or early holometabolan insects. The presence of damage features, together with a range of probable defensive structures (hairs, spines, apical horns, and thick integuments), suggests that an active arms race involving insects and plant reproductive structures was already well established by the early Permian.