Basal Cyclorrhapha in Amber from the Cretaceous and Tertiary(Insecta: Diptera), and Their Relationships: Brachycera in Cretaceous Amber Part IX

ABSTRACT Diverse new basal (aschizan) Cyclorrhapha fossilized in amber are described from the Tertiary and Cretaceous, and their relationships are examined with character-based phylogenetic hypotheses for each family or family group. There are 18 new species in 15 genera (11 of them new) and four families plus the Syrphoidea. Fossils are from the Early Cretaceous of Lebanon, Late Cretaceous of New Jersey (United States) and Alberta (Canada), Eocene of the eastern Baltic coast, and Miocene of the Dominican Republic, but predominantly from the mid-Cretaceous of Myanmar. Stem-group Lonchopteroidea are Alonchoptera lebanica, n. gen., n.sp., and Lonchopterites burmensis, n. sp. Platypezidae include the stem groups Burmapeza radicis, n. gen., n. sp., Canadopeza biacrosticha, n. gen., n. sp., and Calvopeza divergens, n. gen., n. sp. An unnamed Microsania sp. is the first definitive Platypezidae in Baltic amber; Lebanopeza azari, n. gen., n. sp., is a stem group to the Microsaniinae and Melanderomyiinae. Chandleromyia anomala, n. gen., n. sp., is an anomalously derived Platypezinae from the Cretaceous, and two new species of the diverse Recent genus Lindneromyia are in Dominican amber (L. neomedialis and L. dominicana). Fossils of the relict family Ironomyiidae (with 3 living species from eastern Australia) include two stem-group genera with two new species each, all in Burmese amber: Palaeopetia dorsalis and P. terminus, Proironia (n. gen.) gibbera and P. burmitica. All other species of Palaeopetia are compression fossils from the Cretaceous of Asia and Eurasia. For Phoridae, a new defining feature is a stridulatum on the procoxa and profemur in both sexes, occuring in most fossil taxa where observable. New sciadocerines include Eosciadocera pauciseta, n. sp., a very large species in Baltic amber, and two stem groups in Burmese amber, Prophora dimorion, n. gen., n. sp., and a very small, undescribed taxon. Archiphora pria Grimaldi and Cumming in Turonian-aged New Jersey amber is transferred to Hennigophora Brown, based on evidence from a new specimen. Prioriphorinae (not taxonomically treated here) is a paraphyletic, Cretaceous grade to the very diverse, crown-group radiation of Euphorida that occurred in the Cenozoic. Two syrphoids occur in Burmese amber: Prosyrphus thompsoni, n. gen., n. sp. (an apparent stem group to the Syrphidae), and Aschizomyia burmensis, n. gen., n. sp. (with more ambiguous affinities). Several immatures of undetermined family are reported, one a probable phorid larva. No definitive Schizophora are yet known from the Cretaceous.

[1]  Xin-Ran Li,et al.  Old method not old-fashioned: parallelism between wing venation and wing-pad tracheation of cockroaches and a revision of terminology , 2018, Zoomorphology.

[2]  Alexey M. Kozlov,et al.  New data, same story: phylogenomics does not support Syrphoidea (Diptera: Syrphidae, Pipunculidae) , 2018 .

[3]  T. Pape,et al.  Remarkable fly (Diptera) diversity in a patch of Costa Rican cloud forest: Why inventory is a vital science. , 2018, Zootaxa.

[4]  T. Pape,et al.  Comprehensive inventory of true flies (Diptera) at a tropical site , 2018, Communications Biology.

[5]  B. Brown,et al.  Puyehuemyia chandleri, gen. nov., sp. nov. (Diptera: Opetiidae): Remnant of a Cretaceous Biota in Chile , 2018, American Museum Novitates.

[6]  M. Engel,et al.  Wing Tracheation in Chrysopidae and Other Neuropterida (Insecta): A Resolution of the Confusion about Vein Fusion , 2017, American Museum Novitates.

[7]  A. Ross.  Extraordinary Lagerstätten in Amber, with particular reference to the Cretaceous of Burma , 2017 .

[8]  J. Vaňhara,et al.  Molecular phylogeny of flat‐footed flies (Diptera: Platypezidae): main clades supported by new morphological evidence , 2017 .

[9]  D. Grimaldi.  Diverse Orthorrhaphan Flies (Insecta: Diptera: Brachycera) in Amber From the Cretaceous of Myanmar: Brachycera in Cretaceous Amber, Part VII , 2016, Bulletin of the American Museum of Natural History.

[10]  M. Reemer,et al.  Anchored enrichment dataset for true flies (order Diptera) reveals insights into the phylogeny of flower flies (family Syrphidae) , 2016, BMC Evolutionary Biology.

[11]  B. Brown,et al.  Revision of the New World Species of the Millipede-Parasitic Genus Myriophora Brown (Diptera: Phoridae). , 2015, Zootaxa.

[12]  B. Brown,et al.  New morphological characters for classifying Phoridae (Diptera) from the structure of the thorax , 2015 .

[13]  T. Tachi.  Homology of the metapleuron of Cyclorrhapha, with discussion of the paraphyly of Syrphoidea (Diptera: Aschiza) , 2014 .

[14]  M. Engel,et al.  A Diverse Paleobiota in Early Eocene Fushun Amber from China , 2014, Current Biology.

[15]  S. B. Archibald,et al.  Early Eocene big headed flies (Diptera: Pipunculidae) from the Okanagan Highlands, western North America , 2014, The Canadian Entomologist.

[16]  B. Brown.  A New Species of Anevrina from Baltic Amber (Diptera: Phoridae) , 2013 .

[17]  B. Sinclair,et al.  Male terminalia of Diptera (Insecta): a review of evolutionary trends, homology and phylogenetic implications , 2013 .

[18]  D. Grimaldi,et al.  Age constraint on Burmese amber based on U–Pb dating of zircons , 2012 .

[19]  S. Marshall.  Flies: The Natural History and Diversity of Diptera , 2012 .

[20]  H. Nakayama.  Complex asymmetric male genitalia of Anevrina Lioy (Diptera: Phoridae). , 2012, Arthropod structure & development.

[21]  P. Tafforeau,et al.  A new species of the Cretaceous genus Prioriphora (Diptera: Phoridae) in French amber , 2011 .

[22]  H. Schmitz,et al.  Fire-Adaptation in Hypocerides nearcticus Borgmeier and Anabarhynchus hyalipennis hyalipennis Marquart and New Notes about the Australian "Smoke Fly" Microsania australis Collart (Diptera: Phoridae, Therevidae and Platypezidae) , 2011 .

[23]  Markus Friedrich,et al.  Episodic radiations in the fly tree of life , 2011, Proceedings of the National Academy of Sciences.

[24]  C. Labandeira,et al.  Distinguishing Agromyzidae (Diptera) Leaf Mines in the Fossil Record: New Taxa from the Paleogene of North America and Germany and their Evolutionary Implications , 2010, Journal of Paleontology.

[25]  Norm Johnson,et al.  The Fossil Insects of the Green River Shales , 2010 .

[26]  B. Brown.  An enigmatic new genus and species of phorid fly from Baltic amber. , 2009 .

[27]  S. Marshall,et al.  Review of the Nearctic Lonchopteridae (Diptera), including descriptions of three new species , 2008, The Canadian Entomologist.

[28]  F. Gilbert,et al.  Phylogenetic relationships and the larval head of the lower Cyclorrhapha (Diptera) , 2008 .

[29]  D. Mcalpine.  New Extant Species of Ironic Flies (Diptera: Ironomyiidae) with Notes on Ironomyiid Morphology and Relationships , 2008 .

[30]  S. Rojo,et al.  First phylogeny of predatory flower flies (Diptera, Syrphidae, Syrphinae) using mitochondrial COI and nuclear 28S rRNA genes: conflict and congruence with the current tribal classification , 2008 .

[31]  B. Brown.  A further new genus of primitive phorid fly (Diptera: Phoridae) from Baltic Amber and its phylogenetic implications , 2007, Contributions in science.

[32]  Paul M. Choate,et al.  Evolution of the Insects , 2006 .

[33]  F. Gilbert,et al.  Phylogeny of Syrphidae (Diptera) inferred from combined analysis of molecular and morphological characters , 2003 .

[34]  P. Chandler.  The Flat-footed Flies (Diptera: Opetiidae and Platypezidae) of Europe , 2001 .

[35]  D. Yeates,et al.  Phylogeny of the syrphoidea (Diptera) inferred from mtDNA sequences and morphology with particular reference to classification of the pipunculidae (Diptera). , 2000, Molecular phylogenetics and evolution.

[36]  B. Brown.  Re-evaluation of the fossil Phoridae (Diptera) , 1999 .

[37]  D. Grimaldi,et al.  Brachyceran Diptera in Cretaceous ambers and Mesozoic diversification of the Eremoneura. , 1999 .

[38]  R. Macphee,et al.  Age and Paleogeographical Origin of Dominican Amber , 1996, Science.

[39]  R. Zack.  Catalogue of the Fossil Flies of the World (Insecta: Diptera) , 1996 .

[40]  D. Feener,et al.  Efficiency of two mass sampling methods for sampling phorid flies (Diptera: Phoridae) in a tropical biodiversity survey , 1995, Contributions in science.

[41]  E. Pike.  HISTORICAL CHANGES IN INSECT COMMUNITY STRUCTURE AS INDICATED BY HEXAPODS OF UPPER CRETACEOUS ALBERTA (GRASSY LAKE) AMBER , 1994, The Canadian Entomologist.

[42]  P. Chandler.  The oriental and Australasian species of Platypezidae (Diptera) , 1994 .

[43]  R. Disney.  Scuttle Flies: The Phoridae , 1994, Springer Netherlands.

[44]  M. Meyer,et al.  Generic classification of the family Pipunculidae (Diptera) : a cladistic analysis , 1992 .

[45]  B. Brown,et al.  Three new fossil phorid flies diptera phoridae from canadian late cretaceous amber , 1990 .

[46]  A. R. Ennos,et al.  The implications of function on the origin and homologies of the dipterous wing , 1989 .

[47]  J. McAlpine.  Phylogeny and classification of the Muscomorpha , 1989 .

[48]  P. Ferrar.  A guide to the breeding habits and immature stages of Diptera Cyclorrhapha (Part 1: text; Part 2: figures). , 2015 .

[49]  F. Thompson,et al.  Chapter 52. Syrphidae , 1987 .

[50]  H. J. Teskey.  Morphology and terminology - larvae , 1981 .

[51]  J. McAlpine.  Morphology and Terminology- adults , 1981 .

[52]  A. Templeton,et al.  EVOLUTION OF CLONAL DIVERSITY IN THE PARTHENOGENETIC FLY LONCHOPTERA DUBIA , 1980, Evolution; international journal of organic evolution.

[53]  W. Hennig.  Das Hypopygium von Lonchoptera lutea Panzer und die phylogenetischen Verwandtschaftsbeziehungen der Cyclorrhapha (Diptera) , 1976 .

[54]  J. McAlpine.  A FOSSIL IRONOMYIID FLY FROM CANADIAN AMBER (DIPTERA: IRONOMYIIDAE) , 1973, The Canadian Entomologist.

[55]  G. Griffiths.  The phylogenetic classification of diptera Cyclorrhapha : with special reference to the structure of the male postabdomen , 1974 .

[56]  J. McAlpine.  A DETAILED STUDY OF IRONOMYIIDAE (DIPTERA: PHOROIDEA) , 1967, The Canadian Entomologist.

[57]  J. McAlpine,et al.  Systematics of Sciadoceridae and Relatives With Descriptions of Two New Genera and Species From Canadian Amber and Erection of Family Ironomyiidae (Diptera: Phoroidea) , 1966, The Canadian Entomologist.

[58]  E. L. Kessel.  Lindneromyia, a New Genus of Flat-footed Flies, and the Descriptions of Two New Species of Platypezidae from Africa (Diptera) , 1965 .

[59]  F. M. Hull.  A New Genus and Four New Species of Fossil Diptera from Montana and Colorado , 1960 .

[60]  H. Stalker.  ON THE EVOLUTION OF PARTHENOGENESIS IN LONCHOPTERA (DIPTERA) , 1956 .

[61]  F. M. Hull.  A revisional study of the fossil Syrphidae , 1945 .

[62]  J. Malloch.  A Preliminary Classification of Diptera, Exclusive of Pupipara, Based Upon Larval and Pupal Characters, with Keys to Imagines in Certain Families , 1917 .

[63]  Arthur White.  The Diptera-brachycera of Tasmania. Part II , 1915 .

[64]  T. Cockerell,et al.  Fossil insects from Florissant, Colorado , 1910 .

[65]  J. H. Comstock,et al.  The Wings of Insects , 1898, The American Naturalist.

[66]  J. H. Comstock,et al.  A manual for the study of insects, by John Henry Comstock ... and Anna Botsford Comstock ... , 1895 .