Skip to main content Skip to main navigation menu Skip to site footer
Type: Article
Published: 2018-09-19
Page range: 491–510
Abstract views: 95
PDF downloaded: 88

Morphology and life history of an ant parasitoid, Psilocharis afra (Hymenoptera: Eucharitidae)

Department of Entomology, University of California, Riverside, CA 92521
Department of Entomology, University of California, Riverside, CA 92521
Research and Exhibitions Department, Iziko South African Museum, P.O. Box 61, Cape Town, 8000, South Africa 3Department of Biological Sciences, University of Cape Town, Private Bag, Rondebosch, 7701, South Africa
Research and Exhibitions Department, Iziko South African Museum, P.O. Box 61, Cape Town, 8000, South Africa 3Department of Biological Sciences, University of Cape Town, Private Bag, Rondebosch, 7701, South Africa
Hymenoptera Formicidae parasitoid behavior morphology planidium first instar larva


Eucharitidae (Hymenoptera) are specialized ant (Formicidae) parasitoids. As we begin to develop a better understanding of their phylogenetic relationships, it is critical to establish baselines for morphological and biological data. A morphological review and the first report of life history data for Psilocharis afra Heraty is provided based on new material from the Mpumalanga Province of South Africa. Psilocharis Heraty is included in Eucharitinae, but it is unclear whether it is the sister group of all other members of the subfamily, or sister group to Neolosbanus Girault in a monophyletic Psilocharitini, which would in turn be sister group to Eucharitini. The oviposition habits of P. afra differ from those of other Eucharitidae in that eggs are placed among trichomes under bracts at flower bases, instead of either being inserted into cavities formed in plant tissue by an enlarged ovipositor (as in Oraseminae and some Neolosbanus) or inserted into cavities in plant tissue, as in most Eucharitini. The egg and first-instar planidia larva are described, and adult morphology is discussed with reference to Eucharitidae and other parasitoid Hymenoptera.



  1. Ayre, G.L. (1962) Pseudometagea schwarzii (Ashm.) (Eucharitidae: Hymenoptera), a parasite of Lasius niger Emery (Formicidae: Hymenoptera). Canadian Journal of Zoology, 40, 157–164.

    Bouček, Z. (1952) The first revision of the European species of the family Chalcididae (Hymenoptera). Acta Entomologica Musei Nationalis Pragae, 27, 1–108.

    Bucher, G.E. (1948) The anatomy of Monodontomerus dentipes BOH., an entomophagous chalcid. Canadian Journal of Research, 26, 230–281.

    Burks, R.A., Heraty, J.M., Dominguez, C. & Mottern, J.L. (2018) Complex diversity in a mainly tropical group of ant parasitoids: Revision of the Orasema stramineipes species group (Hymenoptera: Chalcidoidea: Eucharitidae). Zootaxa, 4401 (1), 1–107.

    Burks, R.A., Heraty, J.M., Mottern, J., Dominguez, K. & Heacox, S. (2017) Biting the bullet: revisionary notes on the Oraseminae of the Old World (Hymenoptera, Chalcidoidea, Eucharitidae). Journal of Hymenoptera Research, 55, 139–188.

    Chiappini, E. & Mazzone, E. (2000) Differing morphology and ultrastructure of the male copulatory apparatus in species-groups of Anagrus Haliday (Hymenoptera: Mymaridae). Journal of Natural History, 34, 1661–1676.

    Clancy, D.W. (1946) The insect parasites of the Chrysopidae (Neuroptera). University of California Publications in Entomology, 7, 403–496.

    Clausen, C.P. (1940a) The immature stages of Eucharidae (Hymenoptera). Journal of the Washington Academy of Sciences, 30, 504–516.

    Clausen, C.P. (1940b) The oviposition habits of the Eucharidae (Hymenoptera). Proceedings of the Entomological Society of Washington, 12, 504–516.

    Darling, D.C. (1988) Comparative morphology of the labrum in Hymenoptera: the digitate labrum of Perilampidae and Eucharitidae (Chalcidoidea). Canadian Journal of Zoology, 66, 2811–2835.

    Darling, D.C. & Miller, T.D. (1991) Life history and larval morphology of Chrysolampus (Hymenoptera: Chalcidoidea: Chrysolampinae) in western North America. Canadian Journal of Zoology, 69, 2168–2177.

    Darling, D.C. & Roberts, H. (1999) Life history and larval morphology of Monacon (Hymenoptera: Perilampidae), parasitoids of ambrosia beetles (Coleoptera: Platypodidae). Canadian Journal of Zoology, 77, 1768–1782.

    Delvare, G. (1987) Some important morphological features of the Chalcidini (Hymenoptera: Chalcididae) and their implications in the classification of the tribe. In: Gupta, V.K. (Ed.), Advances in Parasitic Hymenoptera Research: Proceedings of the Second Conference on the Taxonomy and Biology of Parasitic Hymenoptera, Gainsville, November 1987. E. J. Brill, Leiden, pp. 25–64.

    Dominichini, G. (1953) Studio sulla morfologia dell'addome degli Hymenoptera Chalcidoidea. Bollettino di Zoologia Agraria e Bachicoltura, 19, 183–298.

    Ernst, A., Mikó, I. & Deans, A.R. (2013) Morphology and function of the ovipositor mechanism in Ceraphronoidea (Hymenoptera, Apocrita). Journal of Hymenoptera Research, 33, 25–61.

    Ferris, G.F. (1940) The myth of the thoracic sternites of insects. Microentomology, 5, 87–90.

    Gibson, G.A.P. (1997) Chapter 2. Morphology and terminology. In: Gibson, G.A.P., Huber, J.T. & Woolley, J.B. (Eds.), Annotated keys to the genera of Nearctic Chalcidoidea (Hymenoptera). National Research Council of Canada Research Press, Ottawa, 794 pp.

    Girault, A.A. (1908) A monographic catalogue of the mymarid genus Alaptus Haliday, with descriptions of three new North American forms and of Alaptus iceryae Riley from type material. Annals of the Entomological Society of America, 1, 179–195.

    Goulet, H. & Huber, J.T. (1993) Hymenoptera of the World: an identification guide to families: Agriculture Canada Research Branch, Ottawa, 668 pp.

    Heraty, J.M. (1994) Classification and evolution of the Oraseminae in the Old World, with revisions of two closely related genera of Eucharitinae (Hymenoptera: Eucharitidae). Life Sciences Contributions, Royal Ontario Museum, 157, 1–174.

    Heraty, J.M. (2000) Phylogenetic relationships of Oraseminae (Hymenoptera: Eucharitidae). Annals of the Entomological Society of America, 93, 374–390.[0374:PROOHE]2.0.CO;2

    Heraty, J.M. (2002) A revision of the genera of Eucharitidae (Hymenoptera: Chalcidoidea) of the World. Memoirs of the American Entomological Institute, 68, 1–359.

    Heraty, J.M. (2003) Three new species of Gollumiella Hedqvist (Hymenoptera: Eucharitidae). Zootaxa, 497 (1), 1–10.

    Heraty, J.M., Burks, B.D., Cruaud, A., Gibson, G., Liljeblad, J., Munro, J., Rasplus, J.-Y., Delvare, G., Janšta, P., Gumovsky, A. V., Huber, J.T., Woolley, J.B., Krogmann, L., Heydon, S., Polaszek, A., Schmidt, S., Darling, D.C., Gates, M.W., Mottern, J.L., Murray, E., Dal Molin, A., Triapitsyn, S., Baur, H., Pinto, J.D., van Noort, S., George, J. & Yoder, M. (2013) A phylogenetic analysis of the megadiverse Chalcidoidea (Hymenoptera). Cladistics, 29, 466–542.

    Heraty, J.M. & Darling, D.C. (1984) Comparative morphology of the planidial larvae of Eucharitidae and Perilampidae (Hymenoptera: Chalcidoidea). Systematic Entomology, 9, 309–328.

    Heraty, J.M. & Darling, D.C. (2007) A new genus and species of Perilampidae (Hymenoptera: Chalcidoidea) with uncertain placement in the family. Journal of the Entomological Society of Ontario, 138, 33–47.

    Heraty, J.M. & Darling, D.C. (2009) Fossil Eucharitidae and Perilampidae (Hymenoptera: Chalcidoidea) from Baltic Amber. Zootaxa, 2306, 1–16.

    Heraty, J.M. & Hawks, D. (1998) Hexamethyldisilazane: A chemical alternative for drying insects. Entomological News, 109, 369–374.

    Heraty, J.M., Hawks, D., Kostecki, J.S. & Carmichael, A.E. (2004) Phylogeny and behaviour of the Gollumiellinae, a new subfamily of the ant-parasitic Eucharitidae (Hymenoptera: Chalcidoidea). Systematic Entomology, 29, 544–559.

    Heraty, J.M. & Murray, E. (2013) The life history of Pseudometagea schwarzii, with a discussion of the evolution of endoparasitism and koinobiosis in Eucharitidae and Perilampidae (Chalcidoidea). Journal of Hymenoptera Research, 35, 1–15.

    Heraty, J.M. & Quicke, D.L.J. (2003) Phylogenetic implications of ovipositor structure in Eucharitidae and Perilampidae (Hymenoptera: Chalcidoidea). Journal of Natural History, 37, 1751–1764.

    Heraty, J.M., Woolley, J.B. & Darling, D.C. (1997) Phylogenetic implications of the mesofurca in Chalcidoidea (Hymenoptera), with an emphasis on Aphelinidae. Systematic Entomology, 22, 45–65.

    Heydon, S. (1997) A review of the world genera of the Trigonoderini, with a revision of the species of North America north of Mexico (Hymenoptera: Pteromalidae). Contributions of the American Entomological Institute, 30, 1–84.

    Johnson, J.B., Miller, T.D., Heraty, J.M. & Merickel, F.W. (1986) Observations on the biology of two species of Orasema (Hymenoptera: Eucharitidae). Proceedings of the Entomological Society of Washington, 88, 542‒549.

    Karlsson, D. & Ronquist, F. (2012) Skeletal morphology of Opius dissitus and Biosteres carbonarius (Hymenoptera: Braconidae), with a discussion of terminology. PLoS One, 7, e32573.

    Kirby, W. & Spence, W. (1826) An Introduction to Entomology or, Elements of the Natural History of Insects. Vol. 3. Longman, Rees, Orme, Brown and Green, London, 732 pp.

    Krenn, H.W. & Pass, G. (1995) Morphological diversity and phylogenetic analysis of wing circulatory organs in insects, part II. Holometabola. Zoology, 98, 147–164.

    Krogmann, L. & Vilhelmsen, L. (2006) Phylogenetic implications of the mesosomal skeleton in Chalcidoidea (Hymenoptera, Apocrita) – tree searches in a jungle of homoplasy. Insect Systematics, 20, 615–674.

    Lachaud, J.P. & Pérez-Lachaud, G. (2012) Diversity of species and behavior of hymenopteran parasitoids of ants: A review. Psyche, 2012, 1–24.

    Laing, J.E. & Heraty, J.M. (1981) The parasite complex of the overwintering population of Epiblema scudderiana (Lepidoptera: Olethreutidae) in Southern Ontario. Proceedings of the Entomological Society of Ontario, 112, 59–67.

    MacGillivray, A.D. (1923) External Insect-Anatomy. Scarab Company, Urbana, viii + 388 pp.

    Martin, J. F. (1916) The thoracic and cervical sclerites of insects. Annals of the Entomological Society of America, 9, 38–83.

    Masner, L. (1979) The variicornis-group of Gryon Haliday (Hymenoptera: Scelionidae). The Canadian Entomologist, 111, 791–805.

    Matsuda, R. (1970) Morphology and evolution of the insect thorax. Memoirs of the Entomological Society of Canada, 102, 1–431.

    Michener, C.D. (1944) Comparative external morphology, phylogeny, and a classification of the bees (Hymenoptera). Bulletin of the American Museum of Natural History, 82, 151–326.

    Michener, C.D. (1956) Hymenoptera. In: Tuxen, S.L. (Ed.), Taxonomist’s Glossary of Genitalia in Insects. S.H Service Agency, Copenhagen, pp. 359.

    Munro, J.B., Heraty, J., Burks, R.A., Hawks, D., Mottern, J.L., Cruaud, A., Rasplus, J.-Y. & Janšta, P. (2011) A molecular phylogeny of the Chalcidoidea (Hymenoptera). PLoS One, 6, e27023.

    Murray, E., Carmichael, A.E. & Heraty, J.M. (2013) Ancient host shifts followed by host conservatism in a group of ant parasitoids. Proceedings of the Royal Society, B, 280, 1–9. [20130495]

    Pinto, J.D. (1992) Novel taxa of Trichogramma from the new world Tropics and Australia (Hymenoptera: Trichogrammatidae). Journal of the New York Entomological Society, 100, 621–633.

    Romero, C.A. & Heraty, J.M. (2010) Ultrastructure of scutellar sensilla in Aphytis melinus (Hymenoptera: Aphelinidae) and morphological variation across Chalcidoidea. Journal of Hymenoptera Research, 19, 159–178.

    Ronquist, F. & Nordlander, G. (1989) Skeletal morphology of an archaic cynipoid, Ibalia rufipes (Hymenoptera: Ibaliidae). Entomologica scandinavica Supplement, 33, 1–60.

    Sanger, C. & King, P.E. (1971) Structure and function of the male genitalia in Nasonia vitripennis (Walker) (Hym.: Pteromalidae). The Entomologist, 104, 137–149.

    Smith, H.S. (1916) The habit of leaf-oviposition among the parasitic Hymenoptera. Psyche, 24, 63–68.

    Snodgrass, R.E. (1910a) The anatomy of the honeybee. U.S. Department of Agriculture, Bureau of Entomology Technical Series, 18, 1–162.

    Snodgrass, R.E. (1910b) The thorax of Hymenoptera. Proceedings of the United States National Museum, 39, 37–91.

    Snodgrass, R.E. (1941) The male genitalia of Hymenoptera. Smithsonian Miscellaneous Collections, 99, 1–86.

    Snodgrass, R.E. (1942) The skeleto-musculature of the honey bee. Smithsonian Miscellaneous Collections, 103, 1–120.

    Snodgrass, R.E. (1957) A revised interpretation of the external reproductive organs of male insects. Smithsonian Miscellaneous Collections, 133, 1–60.

    Torréns, J. (2013) A review of the biology of Eucharitidae (Hymenoptera: Chalcidoidea) from Argentina. Psyche, 2013 (926572), 1–14.

    Torréns, J. & Heraty, J.M. (2013) A new genus of Eucharitidae (Hymenoptera: Chalcidoidea), with notes on life history and immature stages. Zootaxa, 3630 (2), 347–358.

    Torréns, J., Heraty, J.M., Murray, E. & Fidalgo, P. (2016) Biology and phylogenetic studies of Lasiokapala Ashmead (Chalcidoidea: Eucharitidae) with description of a new species from Argentina. Systematic Entomology, 41, 596–606.

    Vilhelmsen, L. (2000a) Before the wasp-waist: comparative anatomy and phylogenetic implications of the skeleto-musculature of the thoraco-abdominal boundary region in basal Hymenoptera (Insecta). Zoomorphology, 119, 185–221.

    Vilhelmsen, L. (2000b) The ovipositor apparatus of basal Hymenoptera (Insecta): phylogenetic implications and functional morphology. Zoologica Scripta, 29, 319–345.

    Vilhelmsen, L., Mikó, I. & Krogmann, L. (2010) Beyond the wasp-waist: structural diversity and phylogenetic significance of the mesosoma in apocritan wasps (Insecta: Hymenoptera). Zoological Journal of the Linnean Society, 159, 22–194.

    Yoder, M.J., Mikó, I., Seltmann, K.C., Bertone, M.A. & Deans, A.R. (2010) A gross anatomy ontology for Hymenoptera. PLoS One, 5, e15991.