Arthropod coprolites and wound reaction in the late Paleozoic climbing fern Hansopteris

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Abstract
Interactions between arthropods and plants have been documented extensively in late Paleozoic trees and ground cover plants, but they have rarely been recorded in late Paleozoic climbers. In this study, we present the second example of coprolites preserved within the plant tissue from the early Permian fossil Lagerstätte Wuda Tuff Flora. The host axis is identified as a phyllophore of the climbing fern Hansopteris uncinatus by the combined evidence of anatomy, morphology, and associated plants. Unlike the first coprolites, which were suggested to be produced by oribatid mites, the culprit of the studied coprolites was likely a myriapod or beetle, indicated by their slightly larger size and the boring behaviour. Furthermore, anomalous parenchymatous cells, sclerenchymatous cells, and metaxylem tracheids have been observed surrounding the tunnel, suggesting responses to traumatic stimulus caused by arthropod damage. This discovery provides an informative example of arthropod herbivory on late Paleozoic climbers and sheds light on how the host plant responded during the early stage of injury.

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