Phylogenetic relationships of nonbiting midges in the subfamily Tanypodinae (Diptera: Chironomidae) inferred from morphology

The nonbiting midge subfamily Tanypodinae represents one of the most diverse lineages of Chironomidae. Despite the wide distribution and high diversity of tanypodine chironomids, the evolutionary history of the subfamily remains poorly understood. Here, we present the first phylogenetic analysis of the subfamily Tanypodinae based on morphological data. Cladistic analyses were conducted using 86 morphological characters from 115 species belonging to 54 tanypodine genera, including the eight currently recognised tribes: Anatopyniini, Clinotanypodini, Coelopyniini, Macropelopiini, Natarsiini, Pentaneurini, Procladiini and Tanypodini. We use characters from fourth-instar larvae, pupae and adults of both sexes. We examine the effects of implied weighting by reanalysing the data with varying values of concavity constant (k). Our analysis supports the monophyly of Tanypodinae with Podonominae as its sister group. All previously proposed tribes are recovered as monophyletic assemblages under a wide range of weighting factors. Under these conditions, the genus Fittkauimyia is the sister group of the remaining Macropelopiini and is erected as a new monobasic tribe, Fittkauimyiini trib. n. The tribe Pentaneurini is recovered as monophyletic with some internal relationships resolved. The genus Paramerina, recovered as sister of Reomyia+Zavrelimyia, is formally synonymised with Zavrelimyia syn.n., based on morphological similarity in all three life stages and treated as a subgenus of the latter. Finally, the recently suggested synonymies of Gressittius and Guassutanypus with Alotanypus and the establishment of the subgenera Conchapelopia (Helopelopia), Macropelopia (Bethbilbeckia), Monopelopia (Cantopelopia), Thienemannimyia (Hayesomyia) and Zavrelimyia (Reomyia and Schineriella) are investigated. Our results support all proposed changes, except for the subgenus-level status of Helopelopia and Cantopelopia. We suggest re-establishment of Helopelopia as a genus, but refrain from promoting genus-level status of Cantopelopia at present because the apparent sister-relationship between Monopelopia+ Nilotanypus likely is due to wing vein reduction caused by miniaturisation.