The snakefly family Mesoraphidiidae (Insecta: Raphidioptera) from the Lower Cretaceous Yixian Formation, China: systematic revision and phylogenetic implications

Mesoraphidiidae is one of the major groups among the Mesozoic Raphidioptera, with 70 described species in 30 genera. However, many mesoraphidiid taxa need to be revised and the monophyly of this family is still questionable, lacking conclusive support from convincing autapomorphies. Here, we present a systematic revision of Mesoraphidiidae from the Lower Cretaceous Yixian Formation of north-eastern China. Nine genera and 12 species are described or re-described based on 27 fossil specimens, including four new genera and five new species, namely Allocotoraphidia gen. nov., Yixianoraphidia gen. nov., Beipiaoraphidia martynovi gen. et sp. nov., Grammoraphidia ponomarenkoi gen. et sp. nov., Proraphidia jepsoni sp. nov., Proraphidia yixianensis sp. nov. and Styporaphidia willmanni sp. nov. Two new combinations are proposed, Yixianoraphidia anomala (Ren, 1997) comb. nov. and Allocotoraphidia myrioneura (Ren, 1997) comb. nov. Proraphidia Martynova, 1947 and Styporaphidia Engel & Ren, 2008 are first recorded from the Lower Cretaceous of China. Phiradia Willmann, 1994 stat. rest. is restored as a valid genus. A preliminary phylogenetic analysis is performed to investigate the relationships among the genera of Mesoraphidiidae based on morphological data. The monophyly of the subfamily Alloraphidiinae and the tribe Nanoraphidiini, respectively, were supported in the present analysis; several genera with three forewing radial crossveins or with subradial crossveins (i.e. Cretinocellia Ponomarenko, 1988, Allocotoraphidia gen. nov., Kezuoraphidia Willmann, 1994 and Xuraphidia Hong, 1992) were assigned to earlier diverging groups in Mesoraphidiidae; however, the monophyly of Mesoraphidiinae was not recovered. The present study sheds light on the evolutionary history of Raphidioptera. http://zoobank.org:pub:610CD366-6328-4F5D-9AD0-71A69866C18B

[1]  S. B. Archibald,et al.  The oldest Inocelliidae (Raphidioptera) from the Eocene of western North America , 2019, The Canadian Entomologist.

[2]  C. Labandeira,et al.  Exploiting Nondietary Resources in Deep Time: Patterns of Oviposition on Mid-Mesozoic Plants from Northeastern China , 2019, International Journal of Plant Sciences.

[3]  Rhythms of Insect Evolution , 2019 .

[4]  C. Labandeira,et al.  Maternal care by Early Cretaceous cockroaches , 2019 .

[5]  U. Aspöck,et al.  New species of the snakefly genus Mongoloraphidia (Raphidioptera: Raphidiidae) from China. , 2018, Zootaxa.

[6]  D. Ren,et al.  A remarkable new genus of the snakefly family Mesoraphidiidae (Insecta: Raphidioptera) from the Lower Cretaceous of China, with description of a new species , 2018, Cretaceous Research.

[7]  A. Lemmon,et al.  Evolution of lacewings and allied orders using anchored phylogenomics (Neuroptera, Megaloptera, Raphidioptera) , 2018 .

[8]  M. Engel,et al.  Phylogeny and Evolution of Neuropterida: Where Have Wings of Lace Taken Us? , 2018, Annual review of entomology.

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

[10]  D. Ren,et al.  Systematic revision of the fossil snakefly family Baissopteridae (Insecta: Raphidioptera) from the Lower Cretaceous of China, with description of a new genus and three new species , 2017 .

[11]  Ivonne J. Garzón-Orduña,et al.  Mitochondrial phylogenomics illuminates the evolutionary history of Neuropterida , 2017, Cladistics : the international journal of the Willi Hennig Society.

[12]  P. Goloboff,et al.  TNT version 1.5, including a full implementation of phylogenetic morphometrics , 2016, Cladistics : the international journal of the Willi Hennig Society.

[13]  Xingyue Liu,et al.  New genera and species of the minute snakeflies (Raphidioptera: Mesoraphidiidae: Nanoraphidiini) from the mid Cretaceous of Myanmar. , 2016, Zootaxa.

[14]  Xingyue Liu,et al.  First record of the fossil snakefly genus Mesoraphidia (Insecta: Raphidioptera: Mesoraphidiidae) from the Middle Jurassic of China, with description of a new species. , 2015, Zootaxa.

[15]  A. Rasnitsyn,et al.  New xyelydid sawflies from the Lower Cretaceous of China , 2015 .

[16]  M. Engel,et al.  Blood-Feeding True Bugs in the Early Cretaceous , 2014, Current Biology.

[17]  D. Ren,et al.  New transitional fossil snakeflies from China illuminate the early evolution of Raphidioptera , 2014, BMC Evolutionary Biology.

[18]  S. B. Archibald,et al.  A revision of the late Eocene snakeflies (Raphidioptera) of the Florissant Formation, Colorado, with special reference to the wing venation of the Raphidiomorpha. , 2014, Zootaxa.

[19]  D. Ren,et al.  A remarkable new genus of basal snakeflies (Insecta: Raphidioptera: Priscaenigmatomorpha) from the Early Cretaceous of China , 2013 .

[20]  Shuo Wang,et al.  New Transitional Fleas from China Highlighting Diversity of Early Cretaceous Ectoparasitic Insects , 2013, Current Biology.

[21]  A. Rasnitsyn,et al.  A new sawfly fossil from the lower Cretaceous of China elucidates antennal evolution in the lower Hymenoptera (Pamphilioidea: Praesiricidae: Archoxyelydinae subfam.n.) , 2013 .

[22]  M. Engel,et al.  Snakefly diversity in Early Cretaceous amber from Spain (Neuropterida, Raphidioptera) , 2012, ZooKeys.

[23]  J. Ansorge,et al.  New snakeflies (Insecta: Raphidioptera) from the Lower Cretaceous of the UK, Spain and Brazil , 2011 .

[24]  E. Peñalver,et al.  A new Early Cretaceous snakefly (Raphidioptera: Mesoraphidiidae) from El Soplao amber (Spain , 2010 .

[25]  Xingyue Liu,et al.  Discovery of Amurinocellia H. Aspöck & U. Aspöck (Raphidioptera: Inocelliidae) in China, with description of two new species , 2009 .

[26]  James E. Jepson,et al.  Raphidioptera (Insecta: Neuropterida) from the Lower Cretaceous Purbeck Limestone Group, Dorset, UK , 2009 .

[27]  Pablo A. Goloboff,et al.  TNT, a free program for phylogenetic analysis , 2008 .

[28]  M. Engel,et al.  Fossil snakeflies from the Early Cretaceous of southern Korea (Raphidioptera: Mesoraphidiidae) , 2006 .

[29]  J. Kukalová-Peck,et al.  Relationships among coleopteran suborders and major endoneopteran lineages: Evidence from hind wing characters , 2004 .

[30]  D. Ren,et al.  Flower-associated brachycera flies as fossil evidence for jurassic angiosperm origins , 1998, Science.

[31]  V. Makarkin,et al.  A new fossil species of snakeflies (Raphidioptera: Mesoraphidiidae) from the Late Cretaceous of Kazakhstan, with notes on Turonian Neuropterida , 2015 .

[32]  F. M. Carpenter Cretaceous Insects From Labrador 2. A New Family of Snake-Flies(Neuroptera: Alloraphidiidae) , 1967 .