Mesozoic fossils and the phylogeny of Tipulomorpha (Insecta: Diptera)

Tipulomorpha are known back to the Triassic but are very rare in the fossil record of that period. The oldest tipuloid genus, Archilimonia Krzeminski & Krzeminska, 2003, is redescribed based on the type and a new species from the Middle Triassic (early Anisian, 242-247 Ma) of the northern Vosges Mountains, NE France. Male terminalia of Triassic Tipulomorpha are described for the first time: a terminal gonostylus divided into a fleshy setose clasper and a lobe in Archilimonia grauvogeliana sp. nov. The systematic position of Archilimonia is investigated based on morphological characters of both extant and extinct taxa of Tipulomorpha and other infraorders. Our cladistic analysis is congruent with the family-level relationships recovered by the most recent phylogenetic studies of Tipulomorpha, and the position of some extinct taxa is discussed. The divided gonostylus is considered a synapomorphy of Tipuloidea except Pediciidae, so in our analyses we place Archilimonia between Pediciidae and other Tipuloidea (Limoniidae + (Tipulidae + Cylindrotomidae), with two other Triassic tipuloid genera (Mabelysia and Metarchilimonia). The family Tipulidae including the Mesozoic genus Tipunia is monophyletic, so Tipulidae has existed since the Late Jurassic.