Aculeata

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

Belongs within: Apocrita.
Contains: Chrysididae, Bethylidae, Euaculeata, Dryinidae.

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 have historically been divided between the Euaculeata and Chrysidoidea but recent phylogenetic analyses indicate that the latter group is paraphyletic. ‘Chrysidoid’ families are parasitoids and retain antennae with the same number of segments in both sexes, in contrast to the sexually dimorphic antennae of Euaculeata (Grimaldi & Engel 2005).

Members of the families Dryinidae and Embolemidae develop attached to leafhoppers of the Auchenorrhyncha, protruding from the host’s abdomen in a bulging sac (thylacium). Embolemidae have the ten-segmented antennae inserted remotely from the mouth on a frontal prominence; females are often brachypterous (Naumann 1991). The Sclerogibbidae are a widespread but not diverse group of small wasps attacking webspinners (Embioptera). They have antennae with 15 to 39 segments inserted beneath a frontal shelf (Naumann 1993). The Plumariidae, another small group found in drier regions of South America and southern Africa, are of unknown natural history. Males have a relatively rich wing venation with well developed accessory veins in the apical membrane whereas females are wingless (Brothers 2011). Plumalexius rasnitsyni is a fossil species from Late Cretaceous New Jersey amber that may be related to the Plumariidae (Brothers 2011).

<==Aculeata [Bethyloidea, Chrysidoidea, Cleptiosa, Vespomorpha]
    |  i. s.: StelopolybiaH79
    |           |--S. angulataH79
    |           |--S. areataH79
    |           |--S. falvipennisH79
    |           |--S. multipictaH79
    |           |--S. myrmecophilaH79
    |           `--S. testaceaH79
    |         Lithoserix williamsiGE05
    |         Lancepyris opertusPK17
    |--BethylonymidaeGE05
    |    |--Bethylonymellus cervicalisR02
    |    |--Bethylonymus sibiricusRJ93
    |    `--Meiagaster Rasnitsyn & Ansorge 2000RM-D00
    |         `--*M. cretaceus Rasnitsyn & Ansorge 2000RM-D00
    `--+--+--ChrysididaeBD17
       |  `--+--BethylidaeBD17
       |     `--+--Plumalexius Brothers 2011 [Plumalexiidae]B11
       |        |    `--*P. rasnitsyni Brothers 2011B11
       |        `--PlumariidaeBD17
       |             |--+--Plumarius Philippi 1873B11
       |             |  |    |--P. gradifronsPK17
       |             |  |    `--P. striaticepsB11
       |             |  `--Myrmecopterina Bischoff 1914B11
       |             `--+--Myrmecopterinella Day 1977B11
       |                `--+--Maplurius Roig-Alsina 1994B11
       |                   |    `--M. spatuliferB11
       |                   `--+--Pluroides Diez, Roig-Alsina & Fidalgo 2010B11
       |                      |    `--P. porteriB11
       |                      `--+--Plumaroides Brothers 1974B11
       |                         |    |--P. andalgalensisB11
       |                         |    |--P. brothersiB11
       |                         |    `--P. tiphlus Diez 2008B11
       |                         `--Mapluroides Diez, Fidalgo & Roig-Alsina 2007B11
       |                              `--M. oglobiniB11
       `--+--EuaculeataBD17
          `--+--SclerogibbidaeBD17
             |    |--Probethylus schwarziR00
             |    `--SclerogibbaN91
             |         |--S. africanaB11
             |         |--S. berlandiB11
             |         `--S. magrettiiB11
             `--+--DryinidaeBD17
                `--EmbolemidaeBD17
                     |--EmbolemusN91
                     |    |--E. brothersiB11
                     |    |--E. collinsiB11
                     |    |--E. confusaB11
                     |    `--E. magnaB11
                     `--Ampulicomorpha succinalisRJ93

*Type species of generic name indicated

References

[BD17] Branstetter, M. G., B. N. Danforth, J. P. Pitts, B. C. Faircloth, P. S. Ward, M. L. Buffington, M. W. Gates, R. R. Kula & S. G. Brady. 2017. Phylogenomic insights into the evolution of stinging wasps and the origins of ants and bees. Current Biology 27: 1019–1025.

[B11] Brothers, D. J. 2011. A new Late Cretaceous family of Hymenoptera, and phylogeny of the Plumariidae and Chrysidoidea (Aculeata). ZooKeys 130: 515–542.

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

[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.

[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).

[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.

[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.

[RM-D00] Rasnitsyn, A. P., & X. Martínez-Delclòs. 2000. Wasps (Insecta: Vespida = Hymenoptera) from the Early Cretaceous of Spain. Acta Geologica Hispanica 35 (1–2): 65–95.

[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.

[R00] Ross, E. S. 2000. Embia. Contributions to the biosystematics of the insect order Embiidina. Occasional Papers of the California Academy of Sciences 149: 1–53, 1–36.

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