Abstract
Westphaloneura magnifica gen. et sp. nov., the first known representative of the stem group Neuropteroidea (= Neuropterida + Coleopterida) is described from the Bashkirian/Moscovian of Northern France, in the new family Westphaloneuridae fam. nov. It is based on two forewings and an incomplete hind wing. Its forewing vein media anterior (MA) being well separated from radius/radius posterior vein (R/RP) is a putative symplesiomorphy, also present in the Coleopterida, justifying the proposed position of the fossil. This fossil is contemporaneous with the oldest known Panorpida, sister group of the Neuropteroidea, also from the same area. This first occurrence provides a new key calibration point for the divergence of two Holometabola lineages and will help time-divergence analyses.
References
- Béthoux, O., Cui, Y.Y., Kondratieff, B., Stark, B. & Ren, D. (2011) At last, a Pennsylvanian stem-stonefly (Plecoptera) discovered. BMC Evolutionary Biology, 11 (248), 1–12. https://doi.org/10.1186/1471-2148-11-248
- Béthoux, O. & Nel, A. (2002) Venation pattern and revision of Orthoptera sensu nov. and sister groups. Phylogeny of Palaeozoic and Mesozoic Orthoptera sensu nov. Zootaxa, 96 (1), 1–88. https://doi.org/10.11646/zootaxa.96.1.1
- Breitkreuz, L.C.V., Winterton, S.L. & Engel, M.S. (2017) Wing tracheation in Chrysopidae and other Neuropterida (Insecta): a resolution of the confusion about vein fusion. American Museum Novitates, 3890, 1–44. https://doi.org/10.1206/3890.1
- Kirejtshuk, A.G. & Nel, A. (2013) Skleroptera, a new order of holometabolous insects (Insecta) from the Carboniferous. Zoosystematica Rossica, 22 (2), 247–257. https://doi.org/10.31610/zsr/2013.22.2.247
- Kirejtshuk, A.G. & Nel, A. (2016) Origin of the Coleoptera and significance of the fossil record. Euroasian Entomological Journal, 15 (1), 66–73.
- Kirejtshuk, A.G. & Nel, A. (2020) Reply to the paper “Beutel et al. 2018 (2020). Is †Skleroptera (†Stephanastus) an order in the stemgroup of Coleopterida (Insecta)?” Insect Systematics & Evolution, 51, 375–383. https://doi.org/10.1163/1876312X-00002206
- Kirejtshuk, A.G., Poschmann, M., Prokop, J., Garrouste, R. & Nel, A. (2014) Evolution of the elytral venation and structural adaptations in the oldest Palaeozoic beetles (Insecta: Coleoptera: Tshekardocoleidae). Journal of Systematic Palaeontology, 12 (5), 575–600. https://doi.org/10.1080/14772019.2013.821530
- Kukalová-Peck, J. (1992) Chapter 6: Fossil history and the evolution of hexapod structures. In: Naumann, I.D. (Ed.), The insects of Australia, a textbook for students and research workers (2nd ed.), 1. Melbourne University Press, Melbourne, CSIRO, pp. 141–179.
- Kukalová-Peck, J. & Willmann, R. (1990) Lower Permian ‘mecopteroid-like’ insects from Central Europe (Endopterygota). Canadian Journal of Earth Sciences, 27 (3), 459–468. https://doi.org/10.1139/e90-041
- Molina-Solís, A., Cleal, C.J., Capel, E. & Cascales-Minana, B. (2024) Vegetation dynamics in the Early–Middle Pennsylvanian coal swamps of the Nord-Pas-de-Calais Coalfield, France. Palaeoworld, 33 (2), 328–340. https://doi.org/10.1016/j.palwor.2023.04.005
- Nel, A., Garrouste, R., Peñalver, E., Hernández-Orúe, A. & Jouault, C. (2022) Discovery of the first blattinopsids of the genus Glaphyrophlebia Handlirsch, 1906 (Paoliida: Blattinopsidae) in the Upper Carboniferous of Southern France and Spain and hypothesis on the diversification of the family. Diversity, 14, 1129. https://doi.org/10.3390/d14121129
- Nel, A., Prokop, J., Nel, P., Grandcolas, P., Huang, D.Y., Roques, P., Guilber, E., Dostál, O. & Szwedo, J. (2012) Traits and evolution of wing venation pattern in paraneopteran insects. Journal of Morphology, 273, 480–506. https://doi.org/10.1002/jmor.11036
- Nel, A., Roques, P., Nel, P., Prokop, J. & Steyer, S. (2007) The earliest holometabolous insect: a “crucial” innovation with delayed success (Insecta: Protomeropina: Protomeropidae). Annales de la Société Entomologique de France, (N.S.), 43 (3), 349–355. https://doi.org/10.1080/00379271.2007.10697531
- Nel, A., Roques, P., Nel, P., Prokin, A.A., Bourgoin, T., Prokop, J., Szwedo, J., Azar, D., Desutter-Grandcolas, L., Wappler, T., Garrouste, R., Coty, D., Huang, D.Y., Engel, M..S. & Kirejtshuk, A.G. (2013) The earliest known holometabolous insects. Nature, 503, 257–261. https://doi.org/10.1038/nature12629
- Peters, R.S., Meusemann, K., Petersen, M., Mayer, C., Wilbrandt, J., Ziesmann, T., Donath, A., Kjer, K.M., Aspöck, U., Aspöck, H., Aberer, A., Stamatakis, A., Friedrich, F., Hünefeld, F., Niehuis, O., Beutel, R.G. & Misof, B. (2014) The evolutionary history of holometabolous insects inferred from transcriptome-based phylogeny and comprehensive morphological data. BMC Evolutionary Biology, 14 (52), 1–33. https://doi.org/10.1186/1471-2148-14-52
- Prokop, J, Fernandes, F.R., Lapeyrie, J. & Nel, A. (2015) Discovery of the first lacewings from the Guadalupian of Lodève Basin in southern France (Neuroptera: Permithonidae). Geobios, 48, 263–270. https://doi.org/10.1016/j.geobios.2015.03.001
- Prokop, J., Nel, A. & Engel, M.S. (2023) Diversity, form, and postembryonic development of Paleozoic insects. Annual Review of Entomology, 68, 401–429. https://doi.org/10.1146/annurev-ento-120220-022637
- Prokop, J., Pecharová, M., Garrouste, R., Beattie, R., Chintauan-Marquier, I.C. & Nel, A. (2017) Redefining the extinct orders Miomoptera and Hypoperlida as stem acercarian insects. BMC Evolutionary Biology, 17 (205), 1–20. https://doi.org/10.1186/s12862-017-1039-3
- Schubnel, T., Perdu, L., Roques, P., Garrouste, R. & Nel, A. (2019) Two new stem-stoneflies discovered in the Pennsylvanian Avion locality, Pas-de-Calais, France (Insecta: ‘Exopterygota’). Alcheringa, 43 (3), 430–435. https://doi.org/10.1080/03115518.2019.1569159
- Vallois, B. & Nel, A. (2023) Earliest possible evidence of insect pollination based on a new species of the Carboniferous medullosalean ‘seed’ genus Pachytesta. Review of Palaeobotany and Palynology, 316, 104948. https://doi.org/10.1016/j.revpalbo.2023.104948
- Vasilikopoulos, A., Misof, B., Meusemann, K., Lieberz, D., Flouri, T., Beutel, R.G., Niehuis, O., Wappler, T., Rust, J., Peters, R.S., Donath, A., Podsiadlowski, L., Mayer, C., Bartel, D., Böhm, A., Liu, S.L., Kapli, P., Greve, C., Jepson, J.E., Liu, X.Y., Zhou, X., Aspöck, H. & Aspöck, U. (2020) An integrative phylogenomic approach to elucidate the evolutionary history and divergence times of Neuropterida (Insecta: Holometabola). BMC Evolutionary Biology, 20 (64), 1–24. https://doi.org/10.1186/s12862-020-01631-6
- Willmann, R. (1989) Evolution und Phylogenetisches System der Mecoptera (Insecta: Holometabola). Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft, 544 (1), 1–153.
- Wolfe, J.M., Daley, A.C., Legg, D.A. & Edgecombe, G.D. (2016) Fossil calibrations for the arthropod tree of life. Earth Science Reviews, 160, 43–110. https://doi.org/10.1016/j.earscirev.2016.06.008