Belongs within: Unicalcarida.
Contains: Ichneumonidae, Braconidae, Proctotrupomorpha, Aculeata, Megalyridae, Ceraphronoidea, Evanioidea, Stephanoidea.
The Vespina are a major clade of Hymenoptera united by ancestral parasitoidism and the loss of thoracic limbs in the larvae. Most members belong to the subclade Apocrita, further 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). The remaining modern Vespina belong to the family Orussidae, external parasitoids of wood-boring beetles that retain a broad attachment between thorax and abdomen. The Jurassic Paroryssidae represent stem-group allies of Orussidae. Fossil Orussidae are known from Late Cretaceous amber (Grimaldi & Engel 2005).
Basal relationships within the Apocrita are less firmly established. Peters et al. (2017) supported a clade of parasitoid forms via molecular analysis, that they named Parasitoida, including the Proctotrupomorpha, Ichneumonoidea and Ceraphronoidea. However, a subsequent molecular analysis by Blaimer et al. (2023) placed the Ichneumonoidea as the basalmost branch in the Apocrita. The Ichneumonoidea are united by fusion of the prepectus to the posterolateral margin of the pronotum, articulation of the first and second metasomal segments by dorsolateral condyles on the adjoining tergal margins, and division of the first metasomal sternum between a strongly sclerotised anterior sclerite and a weakly sclerotised posterior region (Grimaldi & Engel 2005). They also have a short scape, and most species have elongate antennae with numerous segments, a trochantellus, and a flexible metasoma often with a strongly exserted ovipositor (Naumann 1991).
Other members of the Apocrita 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. The Stigmaphronidae are a Mesozoic family with similar fore wing veins to modern Ceraphronoidea (with anterior veins fused into a bar along the wing margin) but a single protibial spur and lack an enlarged second metasomal segment (Grimaldi & Engel 2005).
A bunch of apocrites
Published 17 February 2012
During the late nineteenth century, many women attempted to achieve a ‘wasp waist’, using corsets to tighten their waist to as narrow a diameter as possible. The style was so-called, of course, because of its resemblance to the body of a wasp, with a sharp constriction dividing the body. However, this feature is not universal among wasps: rather, it characterises a distinct clade within the wasps, the Apocrita.
Basal members of the Hymenoptera possess a broad junction between thorax and abdomen like that seen in other insects. In apocritan wasps, the first segment of the abdomen became incorporated into the body of the thorax (where it is referred to as the propodeum) and the characteristic wasp waist developed at the front of the second abdominal segment. Because the major divisions of the body in Apocrita therefore do not correspond directly to the thorax and abdomen of other insects, workers on Apocrita instead refer to the mesosoma and metasoma (or ‘altitrunk’ and ‘gaster’). So narrow is the connection between mesosoma and metasoma, in fact, that members of the Apocrita are incapable of taking solid food: only liquids can pass through the waist. This limitation is believed to have later been significant in the development of the social wasps and ants: because mature ants cannot themselves eat solids, they must feed any solid food they collect to their larvae. The larvae then regurgitate the semi-digested food in a liquid form that the adults can handle. This dependance on their larvae induced the formation of stable colonies. Mature wasps that do not form colonies feed on naturally-occurring liquids such as nectar.
Ancestrally, the Apocrita are a lineage of larval parasites, and the majority of species remain so today. The wide distribution of parasites of wood-boring beetles among basal apocritans, and in their sister group the Orussidae among the non-waisted wasps, suggests that this was probably the original lifestyle for the apocritans (Grimaldi & Engel 2005). Living Apocrita can be divided between five main groups: the Stephanidae, the Aculeata (stinging wasps, including all the social forms such as ants and bees), the Ichneumonoidea (ichneumons and braconids), the Proctotrupomorpha, and the Evaniomorpha (though the monophyly of the latter group is debatable). The Stephanidae are a family of long slender beetle parasites that are most diverse in tropical parts of the world.
The evaniomorphs have been suggested to form a group on the basis of the form of the inner articulation of the coxa (the basal segment) of the middle pair of legs, but the polarity of this feature is debatable (Ronquist 1999). The type superfamily, the Evanioidea, includes a group of families characterised by having the articulation of the metasoma to the mesosoma positioned high up on the propodeum rather than low down as in most other wasps. The hatchet wasps of the Evaniidae have a particularly distinctive body form: the mesosoma is boxy, often almost square in side view; the first segment of the metasoma is developed into a long and narrow petiole; and the remainder of the metasoma is relatively small and hangs off the petiole like the head of the eponymous hatchet. Evaniids are parasites of cockroaches, laying their eggs on the cockroaches’ egg cases.
Females of another evaniomorph family, the Trigonalyidae, lay large numbers of eggs inserted into incisions on a plant leaf. When a piece of leaf containing a trigonalyid egg is eaten by a caterpillar, the egg hatches out and the trigonalyid larva emerges, then burrows into the body of the caterpillar. However, the larva’s target is not the caterpillar itself. Instead, the trigonalyid is looking for the parasitic larva of another wasp that may be inside the caterpillar: it is what is called a hyperparasite (that is, a parasite of a parasite). Trigonalyids are also known as parasites of the larvae of social wasps: when the social wasp feeds its larvae on a caterpillar containing a trigonalyid, the trigonalyid may infect the larva to which it is fed (Grimaldi & Engel 2005).
Systematics of Vespina
<==Vespina [Ephialtitoidea, Euhymenoptera, Evaniales, Evaniomorpha, Parasitica]
|--Apocrita [Clistogastra, Ichneumonides, Parasitoida, Proctotrupii, Terebrantes]PK17
| |--IchneumonoideaBS23
| | |--Praeichneumon [Praeichneumonidae]GE05
| | | `--P. townesiRJ93
| | `--+--IchneumonidaePK17
| | `--+--Eoichneumon Jell & Duncan 1986H02 [EoichneumonidaeGE05]
| | | `--E. duncanae Jell & Duncan 1986RJ93
| | `--BraconidaePK17
| `--+--ProctotrupomorphaBS23
| `--+--+--AculeataBS23
| | `--+--MegalyridaeHR11
| | `--Trigonalidae [Archiglossata, Ichneumomimidae, Trigonaloidea, Trigonalyidae]PK17
| | | i. s.: MimelogonalosN91
| | | Pseudogonalos hahniiPK17
| | | Darbigonalus capitatus Rasnitsyn 1986RJ93
| | | Lycogaster pullata [incl. L. pullata nevadensis]S96
| | | Bareogonalos canadensisS96
| | | Poecilogonalos thwaitesiiRD77
| | | Cretogonalys taimyricus Rasnitsyn 1977P92
| | |--Orthogonalys [Orthogonalinae]BS23
| | | `--O. pulchellaHR11
| | `--TrigonalinaeBS23
| | |--TrigonalysB11
| | | |--T. maculatusRD77
| | | |--T. micanticepsB11
| | | `--T. pervetus Cockerell 1917P92
| | `--TaeniogonalosBS23
| | |--T. gundlachii [incl. Poecilogonalos costalis]S96
| | |--T. maculataB11
| | `--T. venatoriaR70
| `--+--+--StigmaphronidaeGE05
| | | |--Aphrostigmon vitimense Rasnitsyn 1991RJ93
| | | |--Stigmaphron Kozlov 1975P92
| | | | `--*S. orphne Kozlov 1975P92
| | | |--Elasmomorpha Kozlov 1975P92
| | | | `--*E. melpomene Kozlov 1975P92
| | | `--Hippocoon Kozlov 1975P92
| | | `--*H. evadne Kozlov 1975P92
| | `--CeraphronoideaBS23
| `--+--EvanioideaPK17
| `--StephanoideaPK17
`--Orussoidea [Idiogastra]PK17
|--Paroryssidae [Parorysidae]GE05
`--OrussidaeHR11
|--Mesorussus taimyrensisGE05
|--Minyorussus luzziiGE05
|--Guiglia sericataN91
|--Orussobaius wilsoniHR11
|--OrussoniaN91
`--Orussus Latreille 1802L02
|--*O. coronatus Latreille 1802L02
|--O. abietinusPK17
|--O. minutusBD17
|--O. occidentalisHR11
|--O. terminalisBS23
`--O. unicolorPK17
Vespina incertae sedis:
KaratavitidaeR02
EphialtitidaeR02
| i. s.: Stephanogaster magnaGE05
| Sippelipterus liasinus Zessin 1985RJ93
|--Karataus Rasnitsyn 1977 [Symphytopterinae]RM-D00
| |--*K. pedalis Rasnitsyn 1977RM-D00
| `--K. hispanicus Rasnitsyn & Martínez-Delclòs 2000RM-D00
`--EphialtitinaeRM-D00
|--LeptephialtitesRM-D00
|--Cratephialtites kourios (Sharkey in Darling & Sharkey 1990)RM-D00, RJ93 [=Karataus kouriosRM-D00]
|--Ephialtites Meunier 1903RM-D00
| `--*E. jurassicus Meunier 1903RM-D00
|--Montsecephialtites Rasnitsyn & Martínez-Delclòs 2000RM-D00
| `--*M. zherikhini Rasnitsyn & Martínez-Delclòs 2000RM-D00
`--Cretephialtites Rasnitsyn & Ansorge 2000RM-D00
`--*C. pedrerae Rasnitsyn & Ansorge 2000RM-D00
Mesomutilla aptera Zhang 1985RJ93
*Type species of generic name indicated
References
[BS23] Blaimer, B. B., B. F. Santos, A. Cruaud, M. W. Gates, R. R. Kula, I. Mikó, J.-Y. Rasplus, D. R. Smith, E. J. Talamas, S. G. Brady & M. L. Buffington. 2023. Key innovations and the diversification of Hymenoptera. Nature Communications 14: 1212.
[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.
[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.
[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.
[P92] Poinar, G. O., Jr. 1992. Life in Amber. Stanford University Press: Stanford.
[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.
[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.