New transitional fossil snakeflies from China illuminate the early evolution of Raphidioptera

Background: Raphidioptera (snakeflies) is a holometabolous order of the superorder Neuropterida characterized by the narrowly elongate adult prothorax and the long female ovipositor. Mesozoic snakeflies were markedly more diverse than the modern ones are. However, the evolutionary history of Raphidioptera is largely unexplored, as a result of the poorly studied phylogeny among fossil and extant lineages within the order. Results: In this paper, we report a new snakefly family, Juroraphidiidae fam. nov., based on exquisitely preserved fossils, attributed to a new species Juroraphidia longicollum gen. et sp. nov., from the Jiulongshan Formation (Middle Jurassic) in Inner Mongolia, China. The new family is characterized by an unexpected combination of plesiomorphic and apomorphic characters of Raphidioptera. Based on our phylogenetic analysis, Juroraphidiidae fam. nov. together with Raphidiomorpha form a monophyletic clade, which is the sister to Priscaenigmatomorpha. The snakefly affinity of Priscaenigmatomorpha is confirmed and another new family, Chrysoraphidiidae fam. nov., is erected in this suborder. Conclusions: Juroraphidiidae fam. nov. is determined to be a transitional lineage between Priscaenigmatomorpha and Raphidiomorpha. Diversification of higher snakefly taxa had occurred by the Early Jurassic, suggesting that these insects had already had a long but undocumented history by this time.