Fossil wing of stem-hymenopteran Avioxyela gallica, from Nel et al. (2013). Scale bar = 1 mm.

Belongs within: Holometabola.
Contains: Tenthredinoidea, Xyelidae, Pamphilioidea, Unicalcarida.

The Hymenoptera include the wasps and their derivative taxa such as bees and ants. Members of the group are primitively characterised by the presence of hamuli, small hooks on the anterior margin of the hind wing that hook onto the fore wing and hold the wings locked together. The fore wings are the principal flight organs with a concordant increase in the mesothoracic musculature. The prothorax is firmly connected to the mesothorax, and the first abdominal segment is integrated into the mesosoma with its sternite much reduced. The order exhibits haplo-diploid sex determination with females emerging from fertilised eggs and males from unfertilised eggs (Ax 1999). Larvae are diverse in form but in basal forms are caterpillar-like with thoracic legs and abdominal prolegs, the latter lacking crochets.

Molecular phylogenetic analysis of Hymenoptera by Peters et al. (2017) divides hymenopterans between the ectophytophagous Eusymphyta and the endophytophagous and parasitic/carnivorous Unicalcarida. However, Blaimer et al. (2023) supported the former grouping as paraphyletic.

Avioxyela gallica is a stem-hymenopteran represented by two preserved fore wings from the Moscovian (late Carboniferous) of Pas-de-Calais, France. It shares the presence of only a few large large rectangular cells between the main veins with modern Hymenoptera, with RP+MA divided in two parallel veins only, and MP and CuA simple (Nel et al. 2013).

<==Hymenoptera (see below for synonymy)
    |--Avioxyela Nel, Engel et al. in Nel, Roques et al. 2013MW15, NR13 [Avioxyelidae]
    |    `--*A. gallica Nel, Engel et al. in Nel, Roques et al. 2013NR13
Hymenoptera incertae sedis:
  Orisema uchancoiH03
  Ophrynopus schauinslandiG27
  Amalthea Rafinesque 1815BR05
  Giraudia Foerster 1868 non Derbès & Solier in Castagne 1851 (ICBN)BR05
  Dineura virididorsata (Retzius 1783)HP-W05
    |--M. guerinii Lucas 1848E12
    `--M. oraniensis Lucas 1848E12
  Tarpa levaillantii Lucas 1849E12
  Scleroderma ruficornis Lucas 1849E12
  ‘Promachus’ Cresson 1887 nec Loew 1848 nec Stål 1875Z01
  Gigantoxyela quadrifurcataNR13
  Baeosomus Thomson 1891Z93
  Patasson nitensM94
    |--C. abietisF92
    `--C. alashanicaF92
  Lygaeonematus laricisF92
  Hoplocampa Hartig 1837M99
    |--H. chrysorrhaea (Klug 1816)M99
    `--H. fulvicornis (Panzer 1801) [incl. H. rutilicornis (Klug 1816)]M99
  Aprosthema Konow 1899M99
    `--A. melanura (Klug 1814) [incl. A. tarda (Klug 1814)]M99
  Strongylogaster Dahlbom 1835M99
    `--S. multifasciata (Geoffroy 1785) [incl. S. lineata (Christ 1791)]M99
  Dulophanes Konow 1907 [incl. Nesoselandria Rohwer 1910]M99
  Ardis Konow 1886M99
    `--A. pallipes (Serville 1823) [incl. A. brunniventris (Hartig 1837)]M99
  Claremontia Rohwer 1909M99
    |--C. alternipes [=Monophadnoides alternipes]M99
    |--C. confusa [=Monophadnoides confusa]M99
    |--C. tenuicornis [=Monophadnoides tenuicornis]M99
    `--C. waldheimii [=Monophadnoides waldheimii]M99
  Fenusa Leach 1817M99
    `--F. pumila Leach 1817 [incl. F. pusilla (Lepeletier 1823)]M99
  Fenusella Enslin 1912 [incl. Messa Leach 1817 (n. n.)]M99
  Scolioneura Konow 1890M99
    |--S. betuleti (Klug 1916)M99
    `--S. vicina Konow 1894M99
  Empria Lepeletier & Serville 1828M99
    |--E. abdominalisF92
    |--E. candidataF92
    |--E. klugii (Stephens 1835) [incl. E. waldstaetterense Liston 1980]M99
    `--E. pulverataF92
  Harpiphorus Hartig 1837M99
    `--H. lepidus (Klug 1818)M99
  Apethymus Benson 1939M99
    |--A. braccatus (Gmelin 1790)M99
    `--A. serotinus (Müller 1776)M99
  Siobla Cameron 1877M99
    `--S. sturmii (Klug 1817)M99
  Aglaostigma Kirby 1882M99
    |--A. fulvipesF92
    `--A. nebulosum (André 1881)M99
  Elinora Benson 1946 [incl. Cuneala Zirngiebl 1956]M99
    `--E. longipes (Konow 1886) [=Allantus longipes; incl. *Cuneala tricolor Zirngiebl 1956]M99
  Blankia Lacourt 1997M99
    `--*B. koehleri (Klug 1817) [=Allantus koehleri]M99
  Sharliphora Wong 1969M99
    `--S. nigella (Förster 1854) [incl. S. ambigua (Fallén 1808) (preoc.)]M99
  Konowia Brauns 1884M99
    `--K. betulae (Enslin 1911) [incl. K. guntionensis Zombori 1969]M99
  Trachelus Jurine 1807M99
    `--T. troglodytus (Fabricius 1787) [=T. troglodyta]M99
  Iota Saussure 1855S00
  Cephenopsis mirabilis Hong 1985RJ93
  Spategaster blattarumR13
  Poecilosoma pulveratumR13
  Pogonius variegatusR13
  Tachus Jurine 1807KM20
  Christolia Brullé 1846EH19
  Myrapetra scutellarisL89

Hymenoptera [Chalastrogastra, Eusymphyta, Neohymenoptera, Orthandria, Piezata, Siricina, Siricomorpha, Symphyta, Tenthredines, Tenthredinetae, Urocerata, Vespida, Vespidea]

*Type species of generic name indicated


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[BR05] Bouchet, P., & J.-P. Rocroi. 2005. Classification and nomenclator of gastropod families. Malacologia 47 (1–2): 1–397.

[EH19] Énay, R., & M. K. Howarth. 2019. Part L, revised, volume 3B, chapter 7: Systematic descriptions of the Perisphinctoidea. Treatise Online 120: 1–184.

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[G27] Gourlay, E. S. 1927. Notes on the New Zealand wood-wasp Ophrynopus schauinslandi Ashmead. Transactions and Proceedings of the New Zealand Institute 57: 691–693.

[H03] Heads, M. 2003. Ericaceae in Malesia: vicariance biogeography, terrane tectonics and ecology. Telopea 10 (1): 311–449.

[HP-W05] Heitland, W., & H. Pschorn-Walcher. 2005. Biology and parasitoids of the peculiar alder sawfly, Platycampus luridiventris (Fallen) (Insecta, Hymenoptera, Tenthredinidae). Senckenbergiana Biologica 85 (2): 215–231.

[KM20] Kury, A. B., A. C. Mendes, L. Cardoso, M. S. Kury & A. A. Granado. 2020. WCO-Lite: Online world catalogue of harvestmen (Arachnida, Opiliones). Version 1.0—Checklist of all valid nomina in Opiliones with authors and dates of publication up to 2018. Published by the author: Rio de Janeiro.

[L89] Lucas, H. 1889. Note suivante relative à des hyménoptères sociaux de la division des méliponides. Annales de la Société Entomologique de France, 6e série 9: cvii–cviii.

[M99] Magis, N. 1999. Répertoire des mouches à scie reconnues en Belgique et au Grand-Duché de Luxembourg (Hymenoptera: Symphyta). Additions et corrections. Notes Fauniques de Gembloux 36: 85–93.

[M94] May, B. M. 1994. An introduction to the immature stages of Australian Curculionoidea. In: Zimmerman, E. C. Australian Weevils (Coleoptera: Curculionoidea) vol. 2. Brentidae, Eurhynchidae, Apionidae and a chapter on immature stages by Brenda May pp. 365–728. CSIRO Australia.

[MW15] McKenna, D. D., A. L. Wild, K. Kanda, C. L. Bellamy, R. G. Beutel, M. S. Caterino, C. W. Farnum, D. C. Hawks, M. A. Ivie, M. L. Jameson, R. A. B. Leschen, A. E. Marvaldi, J. V. McHugh, A. F. Newton, J. A. Robertson, M. K. Thayer, M. F. Whiting, J. F. Lawrence, A. Ślipiński, D. R. Maddison & B. D. Farrell. 2015. The beetle tree of life reveals that Coleoptera survived end-Permian mass extinction to diversify during the Cretaceous terrestrial revolution. Systematic Entomology 40 (4): 835–880.

[NR13] Nel, A., P. Roques, P. Nel, A. A. Prokin, T. Bourgoin, J. Prokop, J. Szwedo, D. Azar, L. Desutter-Grandcolas, T. Wappler, R. Garrouste, D. Coty, D. Huang, M. S. Engel & A. G. Kirejtshuk. 2013. The earliest known holometabolous insects. Nature 503: 257–261.

[R13] Reuter, O. M. 1913. Lebensgewohnheiten und Instinkte der Insekten bis zum Erwachen der sozialen Instinkte. R. Friedländer & Sohn: Berlin.

[RJ93] Ross, A. J., & E. A. Jarzembowski. 1993. Arthropoda (Hexapoda; Insecta). In: Benton, M. J. (ed.) The Fossil Record 2 pp. 363–426. Chapman & Hall: London.

[S00] Siddiqi, M. R. 2000. Tylenchida: Parasites of plants and insects 2nd ed. CABI Publishing: Wallingford (UK).

[Z02] Zherikhin, V. V. 2002. Insect trace fossils. In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 303–324. Kluwer Academic Publishers: Dordrecht.

[Z93] Zimmerman, E. C. 1993. Australian Weevils (Coleoptera: Curculionoidea) vol. 3. Nanophyidae, Rhynchophoridae, Erirhinidae, Curculionidae: Amycterinae, literature consulted. CSIRO Australia.

[Z01] Zompro, O. 2001. A review of Eurycanthinae: Eurycanthini, with a key to genera, notes on the subfamily and designation of type species. Phasmid Studies 10 (1): 19–23.

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