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. Antennae are often more or less distinctly elbowed at the joint between scape and pedicel. The ovipositor has a fairly plesiomorphic morphology for insects, with ventral valves (stylets) moving along the medial valves while the dorsal valves form an ovipositor sheath.

Hymenopterans exhibit haplo-diploid sex determination with females emerging from fertilised eggs and males from unfertilised eggs. 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) divided 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
‘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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[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.

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[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.

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

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[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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