Theronia atalantae, copyright jgerbracht.

Belongs within: Pimplinae.
Contains: Xanthopimpla, Pimpla.

The Pimplini are a group of ichneumon wasps that develop as endoparasites in lepidopteran pupae. Final instar larvae of the Xanthopimpla and Pimpla groups have a strongly developed epistomal arch and totally reduced hypostoma whereas the hypostoma remains well developed in Theronia and other basal genera (Gauld 1984).

Wasps with fangs on their feet?
Published 26 May 2014
Theronia septentrionalis, photographed by Stephen Cresswell.

The ichneumons are one of the more familiar groups of parasitoid wasps for the general public. The species in the photo above is a member of the Theronia group of ichneumons, which attain a reasonable size by wasp standards (a number of species seem to be in the range of 1.5 centimetres long) and are often brightly coloured in yellow or green. The Theronia group is primarily tropical in distribution, though some species are found in more temperate regions. Authors have differed on whether they treat this group as a single genus or divide it between about half a dozen genera; either option is complicated by the fact that both the group as a whole and some of its constituent restricted genera are doubtfully monophyletic (Gauld et al. 2002). Where their larval hosts are known, many members of the Theronia group are endoparasitic in moth cocoons (including some economically significant pests such as the gypsy moth), though at least some species are not parasites of the moth itself but are hyperparasites of other ichneumon larvae attacking the moth. One (sub)genus, Nomosphecia, includes parasites of vespid wasp larvae (Gauld 1984).

Male Theronia atalantae, photographed by Phil Huntley-Franck.

Bright colours are often a sign of danger in the animal kingdom, and the Theronia group seem to follow that trend. One of the group’s distinctive features is larged, curved claws with an associated spatulate bristle. As noted by Gauld (1984), “When caught they sink their large claws into their captor.” This sounds uncomfortable enough in itself, especially as said claws have a tendency to break and leave their tips embedded in the skin if the wasp is not allowed to remove them in her own time. But there’s more: the inside of the claw bears a fluid-filled cavity, and the act of embedding the claws releases the contents of this cavity into the wound. In other words, the claws seem to function in much the same way as the fangs of a venomous snake.

Or do they? We know that the fluid injected by Theronia into would-be attackers can cause irritation to vertebrate epithelium (Gauld et al. 2002), but we don’t seem to know just what it contains or how it acts. As such, we don’t know how confident we can be that the fluid is indeed effective defensively. Theronia may have poison claws that act like fangs. Or it may just have big sharp claws, and that may be enough.

Systematics of Pimplini

Synapomorphies (from Gauld et al. 2002): Pronotum short and deep; anterior margin of pronotum mediodorsally reflexed and directed backwards as a strong pointed tubercle; submetapleural carina complete, strongly expanded into a lobe anteriorly; propodeal spiracle elongately elliptical, more than 3.5 times as long as wide; intercoxal carinae with posterior end of carina meeting its fellow medially, forming a short but strongly raised medial carina reaching back to rim of foramen; hind leg with fourth tarsomere apically obliquely truncated, so inner side overhangs base of fifth tarsomere; hind tarsus of female with orbicula very slender and bar-like, more than 3.5 times as long as wide; hind tarsal claw with large membranous vesicle in both sexes; hind tarsal claw with long conspicuous spatulate bristle, with apex at rest pressed against pointed apex of claw; body ground colour yellowish; sternite I of female with low rounded swelling centrally; laterotergite IV of female moderately broad, 0.25–0.4 times as wide as long, and strongly sclerotised; metasoma of male with tergites VIII and IX completely fused laterally.

|--Theronia Holmgren 1859ANI11 (see below for synonymy)
| | i. s.: ‘Neotheronia’ donovaniGWB02
| | T. flava Gupta 1962AYI11
| | ‘Pimpla’ laevigataG84 [incl. *Pseudocoenites moravicus Kriechbaumer 1892G84, ANI11]
| | T. lucida (Cameron 1903)AYI11 [=*Poecilopimpla lucidaG84]
| | ‘Neotheronia’ mellosaGWB02
| | T. nigrivertex Gupta 1962AYI11
| | T. pseudozebra Gupta 1962 [=T. (Poecilopimpla) pseudozebra]AYI11
| | ‘Neotheronia’ tacubayaGWB02
| | T. viridicansR70
| | T. zebra (Vollenhoven 1879)AYI11 [=Pimpla zebraAYI11; incl. *Orientotheronia rufescens Morley 1913G84]
| |--T. maculosaGWB02 [=Poecilopimpla maculosaGWB02; incl. T. viridicansG84]
| |--+--T. lurida [=Poecilopimpla lurida]GWB02
| | `--T. maskeliyaeGWB02 [=T. (Poecilopimpla) maskeliyaeAYI11]
| | |--T. m. maskeliyaeAYI11
| | `--T. m. schmiedeknechti Krieger 1906 [=T. (Poecilopimpla) maskeliyae schmiedeknechti]AYI11
| |--+--‘Neotheronia’ concolorGWB02
| | `--+--T. melanocera [=Neotheronia melanocera]GWB02
| | `--‘Neotheronia’ rosaiGWB02
| |--+--‘Neotheronia’ nigrolineataGWB02
| | `--+--Augerella Gupta 1962GWB02, ANI11
| | | |--A. insertaANI11
| | | `--A. orientalis [=Theronia (Augerella) orientalis]AYI11
| | | |--A. o. orientalisAYI11
| | | `--A. o. borneensis (Gupta 1962) [=Theronia (Augerella) orientalis borneensis]AYI11
| | `--+--T. clathrata [=Erythrotheronia clathrata]GWB02
| | | |--T. c. clathrataAYI11
| | | |--T. c. javensis Gupta 1962AYI11
| | | `--T. c. malayensis Gupta 1962AYI11
| | `--Parema Gupta 1962GWB02, ANI11
| | |--*P. nigrobalteata (Cameron 1899)G84, AYI11 [=Theronia nigrobalteataG84]
| | | |--P. n. nigrobalteataAYI11
| | | `--P. n. callida (Tosquinet 1903) (see below for synonymy)AYI11
| | `--P. penetrans [=Theronia (Parema) penetrans; incl. T. fumata, T. papuana]G84
| `--+--‘Neotheronia’ montezumaGWB02
| `--+--T. atalantaeGWB02 [=Ichneumon atalantaeG84; incl. Pimpla flavicansG84, *Theronia flavicansANI11]
| `--T. hilarisGWB02
`--+--‘Theronia’ steindachneriGWB02 (see below for synonymy)
`--+--Nomosphecia Gupta 1962GWB02, ANI11
| |--*N. zebroides (Krieger 1906)G84, AYI11 [=Theronia zebroidesAYI11]
| |--N. elegansGWB02
| |--N. insolensANI11
| | |--N. i. insolensANI11
| | `--N. i. ambonaANI11
| |--N. melanosoma [=Theronia (Nomosphecia) melanosoma]G84
| |--N. pyramida (Gupta 1962) [=Theronia (Nomosphecia) pyramida]AYI11
| |--N. scutellata (Gupta 1962) [=Theronia (Nomosphecia) scutellata]AYI11
| `--N. solisiGWB02
`--+--Epitheronia Gupta 1962GWB02, ANI11
| |--E. fidentiaGWB02
| `--E. tomeus (Gupta 1962) [=Theronia (Epitheronia) tomeus]AYI11
`--+--Xanthopimpla groupGWB02
| |--Lissopimpla Kriechbaumer 1889GWB02, G84 (see below for synonymy)
| | |--*L. albopictaGWB02
| | |--L. atra Girault 1924G24
| | |--‘Pimpla’ basalis [incl. *Trichrus stupenda Tosquinet 1903]G84
| | |--L. excelsaJ02 (see below for synonymy)
| | `--L. scutataG84
| `--XanthopimplaGWB02
`--Pimpla groupGWB02
|--Echthromorpha Holmgren 1868GWB02, G84 (see below for synonymy)
| |--E. agrestoria (see below for synonymy)G84
| |--E. atrataGWB02
| |--*Glyptogastra’ hawaiiensis Ashmead 1900G84
| |--E. intricatoria (see below for synonymy)G84
| `--E. nigricornis [incl. E. fastigata, E. maxima]G84
`--+--Stronglyopsis anomalaGWB02
`--+--+--Alophopimpla Momoi 1966GWB02, G84
| | `--*A. polia Momoi 1966G84
| `--ItoplectisGWB02
| | i. s.: I. clavicornisGWB02
| | I. conquisitor [=Pimpla conquisitor, Scambus conquisitor]B37
| |--I. melanocephalaGWB02
| `--+--I. robertiGWB02
| `--+--I. maculatorGWB02
| `--ApechthisGWB02
| |--A. compunctorP93
| |--A. quadridentataGWB02
| `--A. rufataP93

Echthromorpha Holmgren 1868GWB02, G84 [incl. Allotheronia Ashmead 1900G84, Chrysopimpla Cameron 1899G84, Glyptogastra Ashmead 1900G84, Polyamma Kriechbaumer 1894G84, Polyhamma Dalla Torre 1901G84, Rhynchopimpla Kriechbaumer 1894G84, Stagmopimpla Saussure 1892G84, Syene Snellen van Vollenhoven 1878G84]

Echthromorpha agrestoria [=Ichneumon agrestorius; incl. E. conopleura, Pimpla continua, *Polyamma continua, Echthromorpha diversor, *Stagmopimpla hyalina Saussure 1892, E. notulatoria immaculata, E. agrestoria insidiator, Pimpla interrupta, *Rhynchopimpla interrupta, *Echthromorpha maculipennis Holmgren 1868, E. melioratorius, Cryptus notulatorius, *Syene notulatoria, *Chrysopimpla ornatipes Cameron 1899, Echthromorpha platymischa, E. striata]G84

Echthromorpha intricatoria [=Cryptus intricatorius; incl. *Allotheronia duodecimguttata Ashmead 1900, Echthromorpha excavata]G84

Lissopimpla Kriechbaumer 1889GWB02, G84 [incl. Notiopimpla Vachal 1907G84, Trichrus Tosquinet 1903G84, Xenopimpla Cameron 1898G84]

Lissopimpla excelsaJ02 [=Pimpla excelsaG84; incl. L. decemnotataG84, L. haemorrhoidalisG84, L. octoguttata Kriechbaumer 1889G84, *Notiopimpla priocnemidea Vachal 1907G84, Theronia rufipesG84, Rhyssa semipunctataG84, Lissopimpla semipunctataR35, *Xenopimpla semipunctataG84]

Parema nigrobalteata callida (Tosquinet 1903) [=Theronia callida; incl. Orientotheronia maculipes Morley 1913, Xanthopimpla nigrobalteata Cameron 1902 non T. nigrobalteata Cameron 1899]AYI11

Theronia Holmgren 1859ANI11 [incl. Epimecoideus Ashmead 1900ANI11, Erythrotheronia Cameron 1905ANI11, Neotheronia Krieger 1898ANI11, Orientotheronia Morley 1913ANI11, Poecilopimpla Cameron 1903ANI11, Pseudocoenites Kriechbaumer 1892ANI11, G84]

Theronia’ steindachneriGWB02 [incl. T. antheraeG84, T. claripennisG84, T. dubiaG84, Poecilopimpla fumipennisGWB02, T. fumipennisG84, T. teiaeG84]

*Type species of generic name indicated


[ANI11] Amanda, T. P. O., Y. F. Ng & A. B. Idris. 2011. Corrections on the notes of the group-genus Theronia (Hymenoptera: Ichneumonidae: Pimplinae) of Malaysia. Serangga 16 (1): 51–58.

[AYI11] Amanda, T. P. O., S. Yaakop & A. B. Idris. 2011. A catalogue of the genus-group Theronia (Hymenoptera: Ichneumonidae: Pimplinae) from Sundaland. Serangga 16 (1): 75–89.

[B37] Balduf, W. V. 1937. Bionomic notes on the common bagworm, Thyridopteryx ephemeraeformis Haw., (Lepid., Psychidae) and its insect enemies (Hym., Lepid.). Proceedings of the Entomological Society of Washington 39 (7): 169–184.

[G84] Gauld, I. D. 1984. An Introduction to the Ichneumonidae of Australia. British Museum (Natural History).

[GWB02] Gauld, I. D., D. B. Wahl & G. R. Broad. 2002. The suprageneric groups of the Pimplinae (Hymenoptera: Ichneumonidae): a cladistic re-evaluation and evolutionary biological study. Zoological Journal of the Linnean Society 136: 421–485.

[G24] Girault, A. A. 1924. Lese majeste, new insects and robbery. 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: 182).

[J02] Jensen, J. G. 2002. A record of parasitism of tropical grass webworm Herpetogramma licarsisalis (Walker) (Lepidoptera: Pyralidae) by Lissopimpla excelsa (Costa) (Hymenoptera: Ichneumonidae) in New Zealand. New Zealand Entomologist 25: 93–94.

[P93] Pittaway, A. R. 1993. The Hawkmoths of the Western Palaearctic. Harley Books: Colchester.

[R35] Rayment, T. 1935. A Cluster of Bees: Sixty essays on the life-histories of Australian bees, with specific descriptions of over 100 new species. Endeavour Press: Sydney.

[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.

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