Series: Bulletin of the American Museum of Natural History (AMNH Bulletins) Volume: 239
Sixty-five specimens representing 49 species in 37 genera and 12, possibly 13, families of brachycerous Diptera are described in detail. Some genera are family incertae sedis. They are preserved in Cretaceous ambers from the following areas and ages (abbreviations after each are used to designate the following origins of the ambers): Manitoba and Alberta, Canada (C) (Campanian); central New Jersey (NJ) (Turonian); and Lebanon (L) (Neocomian). All taxa described are new species and most genera are described as new, except where noted.
The new taxa and their origins are the following: Tethepomyia thauma (NJ), an extremely apomorphic fly of probable nematocerous affinities. In Rhagionidae: Paleochrysopilus hirsutus (L), Jersambromyia borodini (NJ), Mesobolbomyia acrai (L); and four additional genera (3 L, 1 NJ) that are described and illustrated but not named because of incomplete preservation. Stratiomyidae: a new specimen of Cretaceogaster pygmaeus Teskey (C) is reported, showing newly observed structures that confirm its extremely primitive position in the family; in addition, in NJ amber an additional primitive genus is described but not named, with affinities in the Pachygastrinae, Chiromyzinae, or Beridinae. Hilarimorphidae: Hilarimorphites superba, H. yeatesi, and H. longimedia, all in NJ amber, and the only fossil hilarimorphids. Scenopinidae(?): Proratites simplex (NJ), probably a primitive (proratine) scenopinid, which would be the only Mesozoic fossil of the family. Asilidae: an incomplete, unnamed specimen in NJ amber, which is one of only two Cretaceous records. The most diverse and numerous brachycerans in Cretaceous ambers are in the Empidoidea, with new taxa as follows. Empidinae: Turonempis styx (NJ), Emplita casei (NJ). Atelestinae: Atelestites senectus (L). Nemedina genus group: Cretodromia glaesa (C); Nemedromia campania (C), N. telescopica (C), N. turonia (NJ); Neoturanius asymmetrus (NJ), N. cretatus (NJ), and N. vetus (NJ, possibly also C); Phaetempis lebanensis (L), which is possibly a very plesiomorphic member of this group.
The Nemedina group today is represented by a single extant species from Hungary. Tachydromiinae: Cretoplatypalpus americanus (C), with Cretoplatypalpus Kovalev previously known from a species in Cenomanian amber from northern Siberia; and Mesoplatypalpus carpenteri (C). Trichopezinae: Apalocnemis canadambris (C), which is the only species studied here belonging to an extant genus, Apalocnemis Philippi (previously known only from extant species widespread in distribution). Microphorinae: Microphorites similis and M. oculeus (L), two additional species of the extinct genus Microphorites Hennig, known only from Lebanese amber; Avenaphora hispida (L); Cretomicrophorus novemundus (NJ), the second species in the extinct genus Cretomicrophorus Negrobov, originally known from Cretaceous amber of Siberia; Archichrysotus incompletus (NJ) and A. manitobus (C), the genus also previously known from Siberian amber. Dolichopodidae: Sympycnites primaevus (L), which is the oldest definitive dolichopodid.
Three new species are described in an unusual new genus, Chimeromyia, known only from Lebanese amber: C. intriguea, C. acuta, and C. reducta. Chimeromyia possesses features of Empidoidea and Cyclorrhapha. The few Cyclorrhapha in Cretaceous ambers are all very plesiomorphic. Platypezidae: Electrosania cretica (NJ), the most plesiomorphic known platypezid. Lebambromyia acrai (L), formally unplaced to family, is a plesiomorphic phoroid closely resembling Ironomyiidae (with one living species in Australia and Tasmania, and one extinct species previously described in Canadian amber). Lonchopteridae: Lonchopterites prisca (L) and Lonchopteromorpha asetocella (L), the only definitive fossils of this small, extant family. Sciadoceridae: Archiphora pria (NJ); and Archisciada lebanensis (L), the oldest fossil of the family and perhaps the most plesiomorphic phoroid.
In addition, two new species are described in the Mesozoic genus Prioriphora McAlpine and Martin, P. luzzii and P. casei (both NJ). This is the best represented brachyceran genus in the Cretaceous, although it might be a paraphyletic taxon. Three cyclorrhaphan larvae of uncertain family identities are described, all in NJ amber; one appears similar to Sciadoceridae. Phylogenetic significance of most of these fossils are discussed, as are certain characters of traditional importance in the higher classification of Brachycera, such as the number of aristal articles. The fossils are placed onto cladograms of the lower Brachycera, the Empidoidea, and basal Cyclorrhapha, and a chronology is proposed of the origins of brachyceran families. The Brachycera apparently originated in the Lower Jurassic, with the Asiloidea not diversifying until the Lower Cretaceous. The Eremoneura (Empidoidea + Cyclorrhapha), as expected, show later diversification, with subfamily-level radiations of empidoids in the Lower to mid-Cretaceous, and the most plesiomorphic families of Cyclorrhapha (e.g., Platypezoidea, Phoroidea, Lonchopteridae) appearing in the Lower to mid-Cretaceous. Origins and radiations of the Schizophora almost certainly are of much more recent origin, in the mid to latest Cretaceous and especially the Cenozoic. The diversity and detailed preservation of these fossils contribute exceptional insight into the early evolution of the Brachycera and the Eremoneura in particular.
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