Holotype of Bethylonymellus cervicalis, from Penney & Jepson (2014).

Belongs within: Vespina.
Contains: Chrysidoidea, Vespidae, Tiphiidae, Pompilidae, Mutillidae, Apoidea, Formicidae.

The Aculeata include the stinging wasps, the clade of Hymenoptera in which the ovipositor has been modified into a venomous sting formed from a pair of sharpened, stiffened gonapophyses, with the egg deposited through an opening at the base of the sting. The fossil Bethylonymidae are included on the basis of their overall resemblance with aculeates, though it is uncertain whether they yet possessed a true sting or not. Bethylonymids are known as compression fossils from the latest Jurassic of central Asia (Grimaldi & Engel 2005). Living aculeates other than the Chrysidoidea are grouped into a clade Euaculeata on the basis of their having differing antennal segment numbers in males and females (13 in males, 12 in females; both sexes having identical antennomere numbers in Chrysidoidea), and having the seventh metasomal tergum largely desclerotised and hidden in females (Grimaldi & Engel 2005). The Euaculeata have commonly been divided between the Apoidea and Vespoidea, the latter being characterised by reduction of the prepectus and often sparse setae on the hypopharynx (Grimaldi & Engel 2015), but molecular phylogenetic analyses indicate that 'Vespoidea' in the broad sense is paraphyletic (Johnson et al. 2013).

Subgroups of the Aculeata include the Scoliidae, a group of wasps characterised by a distinct pattern of corrugations ('pseudoveins') in the distal parts of the wings, which develop as parasitoids of burrowing scarabaeid beetle larvae.

<==Aculeata [Vespomorpha]
    |  i. s.: Stelopolybia H79
    |           |--S. angulata H79
    |           |--S. areata H79
    |           |--S. falvipennis H79
    |           |--S. multipicta H79
    |           |--S. myrmecophila H79
    |           `--S. testacea H79
    |         Lithoserix williamsi GE05
    |--Bethylonymellus R02 [Bethylonymidae GE05]
    |    `--B. cervicalis R02
    `--+--Chrysidoidea GE05
       `--Euaculeata [Melliniores, Mutillariae, Pompiloidea, Scolietae, Scolioidea, Sphegimae, Vespoidea] GE05
            |  i. s.: Falsiformicidae GE05
            |         Meria tripunctata PK17
            |         Colpa sexmaculata PK17
            |--Sierolomorpha [Sierolomorphidae] GE05
            `--+--+--Vespidae JB13
               |  `--+--Rhopalosomatidae JB13
               |     |    |--Rhopalosoma nearcticum HH07
               |     |    |--Mesorhopalosoma GE05
               |     |    |--Propalosoma GE05
               |     |    |--Olixon MC13
               |     |    `--Harpagocryptus australiae R70
               |     `--Scolebythidae HR11
               |          |--Ycaploca evansi GE05
               |          |--Pristapenesia [incl. Dominibythus] GE05
               |          |--Clystopsenella GE05
               |          `--Scolebythus madecassus HR11
               `--+--+--+--Tiphiidae JB13
                  |  |  `--Chyphotes [Chyphotinae] JB13
                  |  |       `--C. mellipes RBV08
                  |  `--Pompilioidea PK17
                  |       |  i. s.: Sapygina decemguttata PK17
                  |       |         Monosapyga clavicornis PK17
                  |       |--+--Aglyptacros sulcatus RBV08
                  |       |  `--Pompilidae JB13
                  |       `--+--Mutillidae JB13
                  |          `--Sapygidae GE05
                  |               |--Fedtschenkiinae GE05
                  |               `--Sapyga Latreille 1802 L02 [Sapyginae GE05]
                  |                    |--*S. quinquepunctata [=Scolia quinquepunctata] L02
                  |                    `--S. pumila RBV08
                  `--+--+--Apoidea JB13
                     |  `--+--Formicidae JB13
                     |     `--Armania [Armaniidae] GE05
                     |          `--A. robusta GE05
                     `--+--Bradynobaenidae JB13
                        |    |--Typhoctes GE05
                        |    `--Apterogyna JB13
                        `--Scoliidae JB13
                             |  i. s.: Radumeris radula MC13
                             |         Crioscolia alcione JB13
                             |--Proscolia [Proscoliinae] GE05
                             `--Scoliinae N91
                                  |--Campsomeris tasmaniensis N91, H49
                                  |--Pseudotrielis N91
                                  |--Australelis N91
                                  |--Trisciloa ferruginea N91
                                  `--Scolia RBV08
                                       |--S. bicincta HH07
                                       |--S. distincta Lucas 1849 E12
                                       |--S. hirta PK17
                                       |--S. hortorum L02
                                       |--S. manilae A71
                                       |--S. ruficornis C81
                                       |--S. varicolor Lucas 1849 E12
                                       `--S. verticalis RBV08

*Type species of generic name indicated


[A71] Askew, R. R. 1971. Parasitic Insects. Heinemann Educational Books: London.

[C81] Caltagirone, L. E. 1981. Landmark examples in classical biological control. Annual Review of Entomology 26: 213–232.

[E12] Evenhuis, N. L. 2012. Publication and dating of the Exploration Scientifique de l'Algérie: Histoire Naturelle des Animaux Articulés (1846–1849) by Pierre Hippolyte Lucas. Zootaxa 3448: 1–61.

[GE05] Grimaldi, D., & M. S. Engel. 2005. Evolution of the Insects. Cambridge University Press: New York.

[H49] Hatch, E. D. 1949. The New Zealand forms of Calochilus R. Br. Transactions and Proceedings of the Royal Society of New Zealand 77 (2): 247–249.

[HR11] Heraty, J., F. Ronquist, J. M. Carpenter, D. Hawks, S. Schulmeister, A. P. Dowling, D. Murray, J. Munro, W. C. Wheeler, N. Schiff & M. Sharkey. 2011. Evolution of the hymenopteran megaradiation. Molecular Phylogenetics and Evolution 60: 73–88.

[HH07] Hines, H. M., J. H. Hunt, T. K. O'Connor, J. J. Gillespie & S. A. Cameron. 2007. Multigene phylogeny reveals eusociality evolved twice in vespid wasps. Proceedings of the National Academy of Sciences of the USA 104 (9): 3295–3299.

[H79] Howse, P. E. 1979. The uniqueness of insect societies: Aspects of defense and integration. In: Larwood, G., & B. R. Rosen (eds) Biology and Systematics of Colonial Organisms pp. 345–374. Academic Press: London.

[JB13] Johnson, B. R., M. L. Borowiec, J. C. Chiu, E. K. Lee, J. Atallah & P. S. Ward. 2013. Phylogenomics resolves evolutionary relationships among ants, bees, and wasps. Current Biology 23: 2058–2062.

[L02] Latreille, P. A. 1802. Histoire Naturelle, générale et particulière des crustacés et des insectes vol. 3. Familles naturelles des genres. F. Dufart: Paris.

[MC13] Majer, J. D., S. K. Callan, K. Edwards, N. R. Gunawardene & C. K. Taylor. 2013. Baseline survey of the terrestrial invertebrate fauna of Barrow Island. Records of the Western Australian Museum Supplement 83: 13–112.

[N91] Naumann, I. D. 1991. Hymenoptera (wasps, bees, ants, sawflies). In: CSIRO. The Insects of Australia: A textbook for students and research workers 2nd ed. vol. 2 pp. 916–1000. Melbourne University Press: Carlton (Victoria).

Penney, D., & J. E. Jepson. 2014. Fossil Insects: An introduction to palaeoentomology. Siri Scientific Press: Manchester.

[PK17] Peters, R. S., L. Krogmann, C. Mayer, A. Donath, S. Gunkel, K. Meusemann, A. Kozlov, L. Podsiadlowski, M. Petersen, R. Lanfear, P. A. Diez, J. Heraty, K. M. Kjer, S. Klopfstein, R. Meier, C. Polidori, T. Schmitt, S. Liu, X. Zhou, T. Wappler, J. Rust, B. Misof & O. Niehuis. 2017. Evolutionary history of the Hymenoptera. Current Biology 27 (7): 1013–1018.

[RBV08] Rabeling, C., J. M. Brown & M. Verhaagh. 2008. Newly discovered sister lineage sheds light on early ant evolution. Proceedings of the National Academy of Sciences of the USA 105 (39): 14913–14917.

[R02] Rasnitsyn, A. P. 2002. Superorder Vespidea Laicharting, 1781. Order Hymenoptera Linné, 1758 (=Vespida Laicharting, 1781). In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 242–254. Kluwer Academic Publishers: Dordrecht.

[R70] Riek, E. F. 1970. Hymenoptera (wasps, bees, ants). In: CSIRO. The Insects of Australia: A textbook for students and research workers pp. 867–959. Melbourne University Press.

Last updated: 25 September 2017.

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