Unraveling the evolutionary history of the snakefly family Inocelliidae (Insecta: Raphidioptera) through integrative phylogenetics

Inocelliidae is one of the two extant families of the holometabolan order
Raphidioptera (snakeflies), with the modern fauna represented
by seven genera and 44 species. The evolutionary history
of the family is little known. Here we present the first phylogenetic and
biogeographic analyses based on a worldwide sampling of taxa and datasets
combined with morphological characters and mitochondrial genomes, aiming
to investigate the intergeneric phylogeny and historical biogeography of
Inocelliidae. The phylogenetic inference from the combined analysis of
morphological and molecular data recovered the sister-group relationship
between a clade of (Negha + Indianoinocellia) +
Sininocellia and a clade of Fibla + the
Inocellia clade (interiorly nested by Amurinocellia and
Parainocellia). Amurinocellia stat. rev. and
Parainocellia stat. rev. et emend. nov. are relegated to
subgeneric status within Inocellia, while a newly erected
subgenus of Inocellia, Epinocellia subgen.
nov., accommodates the former Parainocellia
burmana (U. Aspöck and H. Aspöck, 1968) plus a new species
Inocellia (Epinocellia) weii sp. nov. Further,
the Inocellia crassicornis group constitutes the
nominate subgenus Inocellia stat. nov., but the Inocellia
fulvostigmata group is paraphyletic. Diversification within
Inocelliidae is distinguished by an Eocene divergence leading to extant
genera and a Miocene radiation of species. A biogeographic scenario
depicts how the diverse inocelliid fauna from East Asia could have
originated from western North America via dispersal across the Beringia
during the early Tertiary, and how the Miocene ancestors of
Inocellia could have accomplished long-distance dispersals via
the Tibet‐Himalayan corridor or eastern Palaearctic to western
Palaearctic. Our results shed new light specifically on the evolution of
Inocelliidae and, in general, the Raphidioptera.