Oldest Leaf-Mine Trace Fossil from East Asia and Stoichiometric Footprint: Insight Into Ancient Nutritional Flow in a Plant–Herbivore Interaction

The Late Triassic saw a flourish of plant–arthropod interactions. By the Late Triassic, insects had developed all distinct strategies of herbivory, notably including some of the earliest occurrences of leaf-mining. Herein we describe exceptionally well-preserved leaf-mine trace fossils on a Cladophlebis fern pinnule from the Momonoki Formation, Mine Group, Japan (Middle Carnian), representing the oldest unequivocal leaf-mines from East Asia. The mines all display a distinctive frass trail – a continuous meandering line, which later becomes a broad blotch containing spheroidal particles – demonstrating larval development. The shapes of these mines generally resemble those of nepticuloid moths, although they are absent from extant fern-mining assemblages. Furthermore, elemental analyses by X-ray fluorescence (XRF) reveals that the frass trail comprises phosphate coprolites. The quantitative variations in P, S, and Si between coprolites and leaf veins may reflect physiological processes (consumption, absorption, and excretion) mediated by plant chemicals. Our findings reinforce the idea that leaf-mining had become a pervasive feeding strategy of herbivorous insects by the Late Triassic.