The contribution of the Middle Triassic fossil assemblage of Monte San Giorgio to insect evolution

The Triassic represents a critical period for understanding the turnover of insect fauna from the Paleozoic to the Mesozoic following the end-Permian mass extinctions (EPME); however, fossil deposits from the Early-Middle Triassic are scarce. The exceptionally preserved 239 million-year-old fossil insect fauna recorded at Monte San Giorgio (Switzerland), including 248 fossils representing 15 major insect clades is presented here. Besides the exceptional features, including their small size and excellent preservation, the fossils have importance in the evolutionary history of the group. The taxonomic and ecological diversity recovered, including both freshwater (dragonflies and caddisflies) and terrestrial taxa (true bugs and wasps), demonstrates that complex environments sustained a paleocommunity dominated by monurans (thought not to have survived the EPME), midges, and beetles. Interestingly, a blattodean-like fossil bearing an external ootheca was also found, important for understanding Paleozoic roachoids to extant cockroaches' transition and the evolution of maternal brood care. Moreover, the youngest and first complete specimen of †Permithonidae and the oldest sawfly fossils were discovered. Finally, round-shaped bodies, compatible with seminal capsules or lycophyte spores, were found on the abdomens of several midge-like individuals. If these are spores, non-seed-bearing plants could have been the first entomophilous plants rather than gymnosperms, as recently supposed. Altogether, these fossils contribute substantially to understanding insect evolution and Paleozoic-Mesozoic faunal turnover.