Biting midges (Diptera: Ceratopogonidae) from the Early Cretaceous El Soplao amber (N Spain)

Three new species, Lebanoculicoides excantabris, Archiaustroconops borkenti, and Atriculicoides szadziewskii are described from the Early Cretaceous (early Albian) El Soplao amber deposit (Rábago, Cantabria, northern Spain). Protoculicoides skalskii Szadziewski and Arillo, found in the other Albian Spanish ambers from Peñacerrada I (in Burgos) and San Just (in Teruel), and Austroconops sp., are identified from this new outcrop. The find of a new species of Lebanoculicoides Szadziewski is especially significant since this genus is considered the basalmost known among ceratopogonids. To date, the new species of Atriculicoides Remm is the oldest occurrence for this genus. A general review of the taxonomy and phylogeny of the family Ceratopogonidae, and the palaeoecological significance and palaeogeographic distribution of its basalmost lineages are given. The new data extend knowledge about biting midges during the Early Cretaceous, a key period for understanding the phylogenetic relationships of the ancient members of the family.