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). 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).

<==Aculeata [Bethyloidea, Chrysidoidea, Cleptiosa, 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
    |         Lancepyris opertus PK17
    |--Bethylonymidae GE05
    |    |--Bethylonymellus cervicalis R02
    |    `--Bethylonymus sibiricus RJ93
    `--+--+--Chrysididae BD17
       |  `--+--Bethylidae BD17
       |     `--+--Plumalexius Brothers 2011 [Plumalexiidae] B11
       |        |    `--*P. rasnitsyni Brothers 2011 B11
       |        `--Plumariidae BD17
       |             |--+--Plumarius Philippi 1873 B11
       |             |  |    |--P. gradifrons PK17
       |             |  |    `--P. striaticeps B11
       |             |  `--Myrmecopterina Bischoff 1914 B11
       |             `--+--Myrmecopterinella Day 1977 B11
       |                `--+--Maplurius Roig-Alsina 1994 B11
       |                   |    `--M. spatulifer B11
       |                   `--+--Pluroides Diez, Roig-Alsina & Fidalgo 2010 B11
       |                      |    `--P. porteri B11
       |                      `--+--Plumaroides Brothers 1974 B11
       |                         |    |--P. andalgalensis B11
       |                         |    |--P. brothersi B11
       |                         |    `--P. tiphlus Diez 2008 B11
       |                         `--Mapluroides Diez, Fidalgo & Roig-Alsina 2007 B11
       |                              `--M. oglobini B11
       `--+--Euaculeata BD17
          `--+--Sclerogibbidae BD17
             |    |--Probethylus schwarzi R00
             |    `--Sclerogibba N91
             |         |--S. africana B11
             |         |--S. berlandi B11
             |         `--S. magrettii B11
             `--+--Dryinidae BD17
                `--Embolemidae BD17
                     |--Embolemus N91
                     |    |--E. brothersi B11
                     |    |--E. collinsi B11
                     |    |--E. confusa B11
                     |    `--E. magna B11
                     `--Ampulicomorpha succinalis RJ93

*Type species of generic name indicated


[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 LinnĂ©, 1758 (=Vespida Laicharting, 1781). In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 242–254. Kluwer Academic Publishers: Dordrecht.

[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.

Last updated: 13 March 2021.

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