The phylogenetic relationships of Cretaceous biting midges, with a key to all known genera (Diptera: Ceratopogonidae)

The phylogenetic positions of Cretaceous species of Ceratopogonidae previously placed in the genera Archiculicoides Szadziewski, Protoculicoides Boesel, and Atriculicoides Remm are reappraised in light of synapomorphies. Character states are discussed in detail, supported by new photographs of Protoculicoides depressus Boesel, the description of Protoculicoides revelatus, n. sp., from Burmese amber, and a compilation of previously published illustrations. The recent article by Szadziewski et al. (2016) proposing that Protoculicoides and Atriculicoides are congeneric is shown to be inaccurate. At least three separate lineages are represented by species in these two genera, requiring a new genus, Gerontodacus (type species, G. succineus (Szadziewski)), to include some of them. Archiculicoides, Protoculicoides, Gerontodacus, Adelohelea Borkent and Alautunmyia Borkent remain undetermined to subfamily. As a result of phylogenetic and other taxonomic considerations, the following are new combinations: Gerontodacus krzeminskii (Choufani, Azar, and Nel), Gerontodacus punctus (Borkent), Gerontodacus skalskii (Szadziewski and Arillo), Archiaustroconops andersoni (Szadziewski, Ross, and Gilka), Atriculicoides ciliatus (Borkent), Atriculicoides hispanicus (Szadziewski and Arillo), Atriculicoides sanjusti (Szadziewski and Arillo) and Adelohelea burmitica (Szadziewski and Poinar). The following species are returned to the genera they were assigned to before Szadziewski et al. (2016): Atriculicoides cenomanensis Szadziewski and Schluter, Atriculicoides dasyheleis Szadziewski, Atriculicoides globosus (Boesel), Atriculicoides incompletus Szadziewski and Schluter, Atriculicoides macrophthalmus Remm, Atriculicoides sibiricus Szadziewski, Atriculicoides swinhoei (Cockerell), Atriculicoides szadziewskii Perez-de la Fuente, Delclos, Penalver, and Arillo and Atriculicoides taimyricus Szadziewski. A key is provided to all Cretaceous Ceratopogonidae genera.