Middle-Late Triassic insect radiation revealed by diverse fossils and isotopic ages from China.

The Triassic represented an important period that witnessed the diversification of marine and terrestrial ecosystems. The radiations of terrestrial plants and vertebrates during this period have been widely investigated; however, the Triassic history of insects, the most diverse group of organisms on Earth, remains enigmatic because of the rarity of Early-Middle Triassic fossils. We report new insect fossils from a Ladinian deposit (Tongchuan entomofauna) dated to approximately 238 to 237 million years ago and a Carnian deposit (Karamay entomofauna) in northwestern China, including the earliest definite caddisfly cases (Trichoptera), water boatmen (Hemiptera), diverse polyphagan beetles (Coleoptera), and scorpionflies (Mecoptera). The Tongchuan entomofauna is near the Ladinian-Carnian boundary in age, providing a calibration date for correlation to contemporaneous biotas. Our findings confirm that the clade Holometabola, comprising most of the modern-day insect species, experienced extraordinary diversification in the Middle-Late Triassic. Moreover, our results suggest that the diversification of aquatic insects (a key event of the "Mesozoic Lacustrine Revolution") had already begun by the Middle Triassic, providing new insights into the early evolution of freshwater ecosystems.