Ceraphron sp., copyright Pierre Bornand.

Belongs within: Unicalcarida.
Contains: Proctotrupomorpha, Ichneumonoidea, Aculeata, Megalyridae, Evanioidea.

The Apocrita are a clade of wasps characterised by the development of a narrow waist between the first and second abdominal segments with the first abdominal segment being incorporated into the mesosoma as the propodeum (Grimaldi & Engel 2005). Molecular analysis supports recognition of a major clade of parasitoid forms, including the Proctotrupomorpha, Ichneumonoidea and Ceraphronoidea, as the sister taxon of the remaining members (Peters et al. 2017). The Ceraphronoidea are minute wasps characterised by two protibial spurs, an enlarged second metasomal segment, and fore wings with the costal and subcostal + radial veins fused into a bar running along the anterior margin (Grimaldi & Engel 2005). Other members of the clade include the Trigonalidae, hyperparasitoid wasps within caterpillars or sawfly larvae whose females lay their eggs within incisions in leaves where they are ingested by the primary host when feeding (Naumann 1991).

<==Apocrita [Clistogastra, Ichneumonides, Proctotrupii, Terebrantes] PK17
    |--Parasitoida PK17
    |    |--Proctotrupomorpha PK17
    |    `--+--Ichneumonoidea PK17
    |       `--+--Stigmaphronidae GE05
    |          |    |--Aphrostigmon vitimense Rasnitsyn 1991 RJ93
    |          |    `--Stigmaphron orphne RJ93
    |          `--Ceraphronoidea PK17
    |               |  i. s.: Libanophron astarte PK17
    |               |--Megaspilidae GE05
    |               |    |--Lagynodes Forster 1841 GM79 [Lagynodinae N91]
    |               |    `--Megaspilinae N91
    |               |         |--Megaspilus luscipennis MH11
    |               |         |--Conostigmus lazaros N91, PP19
    |               |         `--Dendrocerus carpenteri N91, M83
    |               `--Ceraphronidae [Calliceratidae] GE05
    |                    |--Lygocerus A71
    |                    |--Aphanogmus A71
    |                    `--Ceraphron Jurine 1807 GM79
    |                         |--C. achilles Dodd 1914 GM79
    |                         |--C. bispinosus MH11
    |                         |--C. javensis Girault 1917 G17
    |                         `--C. sulcatum [=Sparasion (Ceraphron) sulcatum] G20
    `--+--+--Aculeata PK17
       |  `--+--Megalyridae HR11
       |     `--Trigonalidae [Archiglossata, Ichneumomimidae, Trigonaloidea, Trigonalyidae] HR11
       |          |--Mimelogonalos N91
       |          |--Orthogonalys pulchella HR11
       |          |--Pseudogonalos hahnii PK17
       |          |--Darbigonalus capitatus Rasnitsyn 1986 RJ93
       |          |--Lycogaster pullata [incl. L. pullata nevadensis] S96
       |          |--Bareogonalos canadensis S96
       |          |--Trigonalys maculatus RD77
       |          |--Poecilogonalos thwaitesii RD77
       |          `--Taeniogonalos HR11
       |               |--T. gundlachii [incl. Poecilogonalos costalis] S96
       |               `--T. venatoria R70
       `--+--Evanioidea PK17
          `--Stephanoidea H02
               |--Eostephanites Hong 2002 [Eostephanitidae] H02
               |    `--*E. tenuis Hong 2002 H02
               `--Stephanidae HR11
                    |  i. s.: Megischus HR11
                    |           |--M. basalis Aguiar 2005 A05
                    |           `--M. furcatus (Lepeletier & Serville 1825) [incl. M. annulator Brullé 1846] A05
                    |         Protostephanus ashmeadi A05
                    |         Electrostephanus A05
                    |           |--E. brevicornis A05
                    |           `--E. neovenatus A05
                    |         Hemistephanus A05
                    |--Stephanus PK17 [Stephaninae GE05]
                    |    |--S. coronator B35 (see below for synonymy)
                    |    `--S. serrator PK17
                    `--Schlettererinae GE05
                         |--Archaeostephanus corae GE05
                         `--Schlettererius cinctipes N91

Stephanus coronator B35 [=Bracon (Stephanus) coronatus G20, Ichneumon coronatus L02, Pimpla coronator B35]

*Type species of generic name indicated


[A05] Aguiar, A. P. 2005. A new and unusual species of Stephanidae (Hymenoptera), with a discussion on its phylogenetic implications. Journal of Hymenoptera Research 14 (1): 1–6.

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

[B35] Boisduval, J. B. 1835. Voyage de Découvertes de l’Astrolabe. Exécuté par ordre du Roi, pendant les années 1826–1827–1828–1829, sous le commandement de M. J. Dumont d'Urville. Faune entomologique de l'océan Pacifique, avec l'illustration des insectes nouveaux recueillis pendant le voyage vol. 2. Coléoptères et autres ordres. J. Tastu: Paris.

[G17] Girault, A. A. 1917. New Javanese Hymenoptera. Privately published (reprinted Gordh, G., A. S. Menke, E. C. Dahms & J. C. Hall. 1979. The privately printed papers of A. A. Girault. Memoirs of the American Entomological Institute 28: 59–71).

[G20] Goldfuss, G. A. 1820. Handbuch der Naturgeschichte vol. 3. Handbuch der Zoologie pt 1. Johann Leonhard Schrag: Nürnberg.

[GM79] Gordh, G., A. S. Menke, E. C. Dahms & J. C. Hall. 1979. The privately printed papers of A. A. Girault. Memoirs of the American Entomological Institute 28: 1–400.

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

[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.

[H02] Hong Y. 2002. Amber Insect of China. Beijing Scientific and Technological Publishing House.

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

[M83] Martin, N. A. 1983. Miscellaneous observations on a pasture fauna: an annotated species list. DSIR Entomology Division Report 3: 1–98.

[MH11] Munro, J. B., J. M. Heraty, R. A. Burks, D. Hawks, J. Mottern, A. Cruaud, J.-Y. Rasplus & P. Jansta. 2011. A molecular phylogeny of the Chalcidoidea (Hymenoptera). PLoS One 6 (11): e27023.

[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).

[PP19] Palencia, L., E. Peñalver, C. E. Prieto & F. J. Poyato-Ariza. 2019. First fossil harvestmen (Arachnida: Opiliones) from Spain and notes on the fossil record of Opiliones. Palaeontologica Electronica 22.1.5A: 1–18.

[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.

[RD77] Richards, O. W., & R. G. Davies. 1977. Imms' General Textbook of Entomology 10th ed. vol. 2. Classification and Biology. Chapman and Hall: London.

[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.

[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.

[S96] Smith, D. R. 1996. Trigonalyidae (Hymenoptera) in the eastern United States: seasonal flight activity, distribution, hosts. Proceedings of the Entomological Society of Washington 98 (1): 109–118.

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