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Animal Protection Systematic Study

Arthropods Associated with Livestock Dung

About Chalcid Wasps (Chalcidoidea)


Chalcidoid or chalcid wasps are one of the most diverse groups of Hymenoptera (bees, ants, wasps) numerically, structurally, and biologically. About 18,600 valid species have been described in over 1,900 genera world-wide (Noyes 1990). In North America there are over 2,600 described species in 706 genera (Grissell and Schauff 1997a). However, these numbers represent only a fraction of true chalcid diversity, and estimates of 60,000 to 100,000 species world-wide do not seem unreasonable. Structurally and biologically, chalcids are probably as diverse as the rest of the parasitic Hymenoptera put together. They range in size from the smallest insect known, Dicopomorpha echmepterygis Mockford (1997), at about 130 microns (0.13 mm), to over 25 mm, including bizarre as well as beautiful winged and wingless forms. Because of their small size and often extreme fragility, special techniques are required to collect and preserve specimens for study. A general overview of these techniques for insects and mites, Collecting and Preserving Insects and Mites: Techniques and Tools, is available on the USDA - SEL site. The most comprehensive published review of methods to collect, preserve and mount chalcid wasps is by Noyes (1982).

Literature on chalcid wasps is vast. Probably the two most important publications for any serious student of chalcidology in North America are: Annotated Keys to the Genera of Nearctic Chalcidoidea (Hymenoptera) by Gibson et al. (1997), and the now significantly outdated but still essential section on Chalcidoidea in Catalog of Hymenoptera in America north of Mexico by Gordh et al. (1979). The first publication provides individuals with the means to identify the families and genera of chalcids known from North America and, for each genus, gives further information on keys to species, number of species known or estimated, host range, and distribution. The second publication provides a list of the chalcid taxa then known from North America, including complete nomenclatural history and references to papers providing information on the taxonomy, biology and morphology of the taxa. Other important references to the taxonomic literature, with emphasis on the North American fauna, are given in the sections below.

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Economic Importance

Although often very beautiful because of their unusual structures and striking colors or metallic hues, chalcids are also very tiny and unobtrusive, not biting or stinging humans, living in their domiciles, or carting off their food. Because of this, even though they are out there in "the billions and billions" (with apologies to Carl Sagan), they remain largely unnoticed and unknown to all but the specialist, and under-appreciated for their role in the environment and benefit to man [seeLaSalle and Gauld (1992) and LaSalle (1993)]. A few chalcids are harmful because they are phytophagous (particularly seed-feeding Eurytomidae) or are hyperparasites, but the large majority are beneficial because they are primary parasites of other insects and arachnids. Consequently, they help regulate insect populations naturally and are one of the most important groups of insects for biological control of pest species. Chalcids have been involved in about two-thirds of biocontrol programs involving Hymenoptera and about one-third of all biocontrol programs in which partial or complete economic control of an insect pest was achieved (Greathead 1986).

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Biological Diversity

The most comprehensive review of chalcid biology and ecology is by Bendel-Janssen (1977). Grissell and Schauff (1997a) state that the host range of chalcids is thought to exceed that of all other insect groups except for the order Diptera. Members are known to attack hosts from 13 orders of insects (Coleoptera, Diptera, Heteroptera, Homoptera, Hymenoptera, Lepidoptera, Neuroptera, Odonata, Orthoptera, Psocoptera, Siphonaptera, Strepsiptera, and Thysanoptera), as well as egg sacs of spiders (Araneae), ticks and gall-forming mites (Acari), cocoons of pseudoscorpions (Pseudoscorpiones), and gall-forming Anguinidae (Nematoda). In addition, phytophagous chalcids are known from six families: Agaonidae, Eulophidae, Eurytomidae, Pteromalidae, Tanaostigmatidae and Torymidae. The Agaonidae and Tanaostigmatidae are thought to be entirely phytophagous and are gall formers. The Agaonidae are entirely responsible for pollinating fig (Ficus) trees. The other known phytophagous chalcids are also gall formers, or are inquilines in galls produced by other insects, or feed in seeds. Possibly as a consequence of their host diversity, chalcids demonstrate all but 2 of 15 feeding types that have been defined for insects, including parasitism of all host life stages from egg to adult, as internal or external parasites, as primary or hyperparasites, and with their eggs laid in, on or away from the host (Grissell and Schauff 1997a). Some chalcids of the family Aphelinidae even parasitize the opposite sex of their own species (Woolley 1997)!

Like other Hymenoptera, most chalcids have a haploid-diploid mechanism of sex determination in which fertilized (diploid) eggs normally develop into females and unfertilized (haploid) eggs normally develop into males (arrhentokous development). However, males are unknown or are very rare for some species and in these species females produce females asexually (thelytokous development). This latter condition appears often to be the result of the presence of maternally inherited cytoplasmic microorganisms, bacteria in the genus Wolbachia. Heat or such antibiotics as tetracycline can kill the bacterium and turn a thelytokous strain into an arrhentokous strain (Stouthamer et al. 1990). It has also been shown that reproductive isolation between closely related species is at least sometimes the result of the presence of Wolbachia or of the species possessing different bacterial 'types'. Viable hybrid offspring can be produced if the species are 'cured' of their endosymbiotic bacterium (Breeuwer and Werren 1993, Breeuwer and Werren 1995, Perrotminnot et al. 1996).

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Because of their small size and structural diversity it is difficult to give easily visible and reliable features that will differentiate all chalcids from all other insects. Those chalcids that are fully winged are like other Hymenoptera in having two sets of membranous wings, with the forewings much larger than the hind wings. More importantly, the forewing does not have any areas (cells) entirely enclosed by veins and there is at most a single vein complex along the leading edge of the wing that normally branches apically [fig.]. Though presence of a submarginal, marginal, postmarginal, and stigmal vein is characteristic of most chalcids, some chalcids have the wing venation even more reduced and a very few lack veins altogether. Further, some other parasitic Hymenoptera, most notably the family Scelionidae (Platygastroidea), have a very similar venation and therefore are often confused with chalcids. However, in platygastroids the posterolateral angle of the pronotum extends back and touches the more or less oval sclerite (tegula) that covers the base of the forewing. In most chalcids the pronotum is more or less distinctly separate from the forewing because there is an intervening sclerite, the prepectus, between the pronotum and mesopleuron [SEM]. Although often difficult to see, chalcids also have a unique placement of the mesothoracic spiracle: it is located between the pronotum and exposed lateral edge of the mesoscutum, often above the anterodorsal angle of the prepectus but at least separate from the anterodorsal margin of the mesopleuron (mesepisternum) [SEM]. Other parasitic Hymenoptera have the spiracle lower down, either between the posterior margin of the pronotum and the anterior margin of the mesopleuron or on the pronotum itself in this same relative position. Almost all chalcids have ridge-like, longitudinal sensilla on the flagellum of the antenna [SEM] and, usually much more conspicuous, many chalcids have a distinct metallic coloration, which almost no platygastroids have.

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Monophyly and Relationships

Gibson (1986a) proposed that Chalcidoidea, including the family Mymaridae, is monophyletic based on three shared derived features (synapomorphies) that are listed above under Recognition as distinguishing features:

He also proposed that Chalcidoidea was the sister group of the family Mymarommatidae based on three other features shared by the two taxa:

Although Gibson (1986a) further proposed that the family Mymaridae is monophyletic based on at least three synapomorphies, he knew of no synapomorphic features for the rest of the Chalcidoidea and therefore could not demonstrate that Mymaridae was the sister group of all other members of the superfamily.

In 1994 Quicke et al. showed that chalcids other than mymarids (and mymarommatids) have a structure of the female ovipositor that differs from all other Hymenoptera: the fused second valvulae have asymmetrical dorsolateral portions that overlap medially to a greater or lesser extent [SEM]. Further, near the base of the second valvulae is a transversely striated band of notal membrane, the laminated bridge. Quicke et al. proposed that both the unique structure of the second valvulae and the laminated bridge were synapomorphies that united all chalcids except mymarids and demonstrated that Mymaridae was the sister group of all other Chalcidoidea.

Heraty et al. (1997) also showed that Chalcidoidea, excluding Mymarommatidae, have a unique structure of another mesothoracic muscle, the furcal-laterophragmal muscle (unlike other Hymenoptera the muscle is attached along the entire length of the laterophragmal apodeme). This is additional evidence for exclusion of the Mymarommatidae as the sister group of Chalcidoidea. They further showed that structure of the furcal-laterophragmal muscle was somewhat different in mymarids than in other chalcids. Different hypotheses for evolution of the particular mymarid structure are possible, but the structure could be interpreted as an intermediate condition in a single transformation series between that of the plesiomorphic condition possessed by mymarommatids and the most apomorphic condition possessed by most other chalcids. If so, this is additional evidence that Mymaridae is the sister group of all other Chalcidoidea.

Finally, Basibuyuk and Quicke (1995) found features of the antennal cleaning organ of the front leg that might be unique for chalcids. They stated that most chalcids have a protibial comb [SEM] and that mymarids have "an indication of this structure". Only ichneumonoids were said to have somewhat similar setae of unknown function. Basibuyuk and Quicke also stated that the setae of the basitarsal comb [SEM] are uniquely structured in Chalcidoidea. The setae were said to be "distinctly flattened" in chalcids and cylindrical in the rest of the Order. However, the state in mymarommatids was not stated explicitly and the scanning electron micrographs included in the paper do not illustrate the difference between flat versus cylindrical. These features need to be examined more comprehensively throughout the chalcids to determine whether different states might indicate phylogenetic relationships within the superfamily.

Though recent studies appear to support hypotheses that Mymarommatidae is the sister group of the Chalcidoidea and that Mymaridae is the sister group of all other chalcids, other relationships are extremely tentative. This includes relationships between Mymarommatidae + Chalcidoidea and other apocritan Hymenoptera and within the superfamily itself. Rasnitsyn (1988) proposed that Chalcidoidea is the sister group of the Platygastroidea (Platygastridae + Scelionidae), based primarily on a similar loss of wing venation. A Chalcidoidea + Platygastroidea relationship was also suggested by the results of recent molecular analysis of the mitochondrial 16S rRNA gene by Dowton et al. (1997).

A sister group relationship between (Chalcidoidea + Mymarommatidae) + Platygastroidea should be considered as no more than a 'testable hypothesis' at present. Some critical taxa, such as Austroniidae, were not included in the molecular analysis and have not been dissected to determine states of the several internal features that are used to define monophyly and relationships of the Chalcidoidea. Also, a (Chalcidoidea + Mymarommatidae) + Platygastroidea relationship appears unlikely based on some plesiomorphic features retained by Mymarommatidae + Chalcidoidea, unless this clade represents a relatively early divergence in the evolution of apocritan Hymenoptera. Members of the family Mymarommatidae first appear in the fossil record in the Upper Cretaceous, Galoromma bezonnaisensis being described by Schlüter (1978) from French amber of Cenomanium age (about 100 mybp). An undetermined genus of 'Mymaridae' was also reported from Lebanese amber (about 125 mybp), though it is quite possible that this is also a mymarommatid. Yoshimoto (1975) described 12 chalcid species in three extant families, Mymaridae, Trichogrammatidae and Tetracampidae, from Canadian Cretaceous amber (about 78 mybp), though the oldest specimens that can be assigned unequivocally to the Chalcidoidea appear in Agapa and Jantardkah Siberian amber (about 90 mybp) (Rasnitsyn and Kulicka 1990). Although relatively old, these ages do not necessarily support an early divergence because several other groups of apocritan Hymenoptera appear even earlier in the fossil record, beginning in the Jurassic.

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Family Classification

The extreme structural diversity of chalcids is reflected partly in the relatively large number of recognized families and genera, and in an historical instability of familial level classification. Boucek (1988b) provides a comprehensive review of the history of chalcid higher classification. Since as late as 1950 anywhere from 9 to 24 families have been recognized, though recently a consensus has been forming around 20 or 21 families, with remaining controversy as to whether the Mymarommatidae should be included in Chalcidoidea or excluded as its own superfamily. The more recent studies of Quicke et al. (1994) and Heraty et al. (1997) support exclusion of mymarommatids.

The 20 generally accepted families of Chalcidoidea are listed under Recognized Families and Major Taxonomic Resources. Monophyly of many of the families remain in doubt because they often seem to intergrade into each other, with 'family level' features sometimes working for only one sex, not being possessed by all members of the family, or being possessed by some members of other families. Consequently, at least some families appear to be more 'taxa of convenience' than monophyletic evolutionary lineages and some diverse families such as the Pteromalidae and Aphelinidae are likely paraphyletic relative to some of the smaller families, and perhaps even polyphyletic. As one might suspect, membership of some families remains controversial. Examples include whether Chrysolampinae should be classified in Pteromalidae or Perilampidae, the Mongolocampinae in Tetracampidae or Aphelinidae, the Eriaporinae in Aphelinidae or Elasmidae, the Akapalinae, Ecthrodapinae and Philomidinae in Eucharitidae or other families (Pteromalidae, Torymidae, Perilampidae), and the Epichrysomalline, Sycoecinae, Otitesellinae, Sycophaginae and Sycoryctinae in Agaonidae or some other families (Pteromalidae, Torymidae). There are even current uncertainties about the proper family placement of some genera, such as Cynipencyrtus Ishii (Tanaostigmatidae or ?), Cales Howard (Aphelinidae or ?), and Idiporous LaSalle and Polaszek (Pteromalidae or ?). Is there any wonder why chalcid wasps remain such a fascinating group of study!

Below are listed some of the major studies that have investigated monophyly and relationships of chalcid families through explicit character state analysis; references are given under the primary family or families investigated though the papers often include discussion of other families.

Shaffee and Rizvi (1990), Heraty et al. (1997), LaSalle et al. (1997)
    Chalcididae and Leucospidae
Wijesekara 1997
    Encyrtidae, Eupelmidae, and Tanaostigmatidae
LaSalle and Noyes (1985), Gibson (1986b), LaSalle (1987), Gibson (1989)
    Eucharitidae and Perilampidae
Heraty and Darling (1984), Darling (1988), Heraty (1989), Darling and Miller (1990), Darling (1992), Heraty (1994)
Gibson (1986a)
Woolley (1988)
Grissell (1995)

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Recognized Families and Major Taxonomic Resources

Listed below in alphabetical order are the 20 families of Chalcidoidea that are generally accepted. Obviously, the relatively few references listed as resources are not intended to be exhaustive. Individuals interested in the North American fauna should refer to the references listed under each family in Gibson et al. (1997). Individuals interested in fauna from other regions should also check references included in works listed under Regional Keys to Families. The references listed below are primarily family-level studies or faunal studies from regions other than North America, but only English language publications.

Ramirez (1970), Ramirez (1974), Wiebes (1982)
Hayat and Verma (1980), Hayat (1983), Yasnosh (1976), Yasnosh (1983), Hayat (1994)
Delvare and Boucek (1992), Wijesekara (1997), Narendran (1989)
Burks (1965)
Tachikawa (1963), Trjapitzin (1973a), Trjapitzin (1973b), Trjapitzin and Gordh (1978a), Trjapitzin and Gordh (1978b), Prinsloo and Annecke (1979), Noyes (1980), Tachikawa (1981), Noyes and Hayat (1984), Noyes (1988), Dahms and Gordh (1997)
Heraty (1985), Heraty (1994)
Graham (1987), Graham (1991)
Gibson (1989), Gibson (1995)
Burks (1971), Stage and Snelling (1986), Zerova (1988), Narendran (1994)
Boucek (1974)
Annecke and Doutt (1961), Subba Rao and Hayat (1983), Schauff (1984), Huber (1986), Viggiani (1988), Noyes and Valentine (1989a), Yoshimoto (1990)
Boucek (1986), Hanson (1992)
Boucek (1978), Darling 1996
Graham (1969), Boucek and Rasplus (1991)
Boucek and Noyes (1987)
Woolley (1986), Woolley (1988)
LaSalle (1987)
Boucek and Askew (1968), Sugonjaev (1971)
Narendran (1994), Grissell (1995)
Doutt and Viggiani (1968), Yousuf and Shafee (1986a), Yousuf and Shafee (1986b), Yousuf and Shafee (1987)

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Regional Keys to Families

It is extremely difficult to briefly characterize or present a key to chalcid families on a world basis, particularly with current uncertainty and controversy over membership of some families, as discussed in Family Classification. The only world key to families that attempts to include exceptional forms is by Gibson (1993), but because of this the key is quite complicated and it is recommended that individuals normally try to identify families using relevant regional keys. For North America, an excellent pictorial key to families and common subfamilies was given by Grissell and Schauff (1997b); a more traditional dichotomous key was also given by Grissell and Schauff (1997a). The following publications that include keys to chalcid families are listed in chronological order; those with an asterisk also include keys to genera.

      Grissell and Schauff (1997a) in * Gibson et al. (1997) - Nearctic region
      Grissell and Schauff (1997b) - Nearctic region
   * Hanson and LaSalle (1995) - Costa Rica
      Naumann (1991) - Australia
   * Noyes and Valentine (1989b) - New Zealand (plus keys to genera of 12 of 16 families)
      Delvare and Aberlenc (1989) - tropical Africa and America
   * Boucek (1988a) - Australasian region (plus keys to genera of 14 of 21 families)
      Gauld and Bolton (1988) - Britain
   * Subba Rao and Hayat (1985) - Oriental region
      Yoshimoto (1984) - Canada
      Prinsloo (1980) - Ethiopian region
   * Medvedev (1978) - European USSR
   * Alayo and Hernandez (1978) - Cuba
      Riek (1970) - Australia
   * Peck, Boucek, and Hoffer (1964) - Europe
      Graham (1969) - Europe (plus key to genera of Pteromalidae)
   * Nikolskaya (1952) - European USSR

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Catalogs are one of the most important resources for taxonomists and biologists alike because they are the means by which all information published on species from 1758 to the present is or can be organized. Catalogs list the described taxa known from a particular region, provide the currently accepted valid name of the taxon, list the synonyms of those valid names, and usually provide additional information and/or references to information on the distribution and biology of the taxa. Listed below are some of the more recent catalogs for the superfamily; catalogs for individual families or genera are not listed.

    Canada and United States
Gordh et al. (1979), Peck (1963)
    Central and South America
De Santis (1967), De Santis (1979), De Santis (1980), De Santis (1983), De Santis (1989), De Santis and Fidalgo (1994)
Alayo (1970)
Subba Rao and Hayat (1986)
Baltazar (1966)

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Alayo D. Pastor 1970.
Catalogo del los Himenopteros de Cuba. Editorial Pueblo y Educacion. 218 pp.
Alayo D. Pastor and L.R. Hernandez. 1978.
Introduccion al estudio de los himenopteros de Cuba: Superfamilia Chalcidoidea. Academia de Ciencias de Cuba, La Habana. 105 pp.
Annecke, D.P. and R.L. Doutt. 1961.
The genera of the Mymaridae (Hymenoptera: Chalcidoidea). Republic of South Africa, Department of Agricultural Technical Services, Entomology Memoirs 5. 71 pp.
Baltazar, C.R. 1966.
A catalogue of Philippine Hymenoptera (with a bibliography, 1758-1963). Pacific Insects Monograph: 8: 1-488.
Basibuyuk, H.H. and Quicke, D.L.J. 1995.
Morphology of the antenna cleaner in the Hymenoptera with particular reference to non-aculeate families (Insecta). Zoologica Scripta 24: 157-177.
Bendel-Janssen, M. 1977.
Zur Biologie Okologie und Ethlogie der Chalcidoidea. Mitteilungen aus der Biologischen Bundesanstalt für Land- und Forstwirtshaft: 176: 1-163.
Boucek, Z. 1974.
A revision of the Leucospidae (Hymenoptera: Chalcidoidea) of the world. Bulletin of the British Museum (Natural History) (Entomology), Supplement 23. 241 pp.
Boucek, Z. 1978.
A generic key to Perilampinae (Hymenoptera: Chalcidoidea), with a revision of Krombeinius n.gen. and Euperilampus Walker. Bulletin of Entomological Research 76: 393-407.
Boucek, Z. 1986.
Taxonomic studies of chalcidoid wasps (Hymenoptera) associated with gall midges (Diptera: Cecidomyiidae) on mango trees. Entomologica Scandinavica 9: 299-307.
Boucek, Z. 1988a.
Australasian Chalcidoidea (Hymenoptera): A biosystematic revision of genera of fourteen families, with a reclassification of species. C.A.B. International, Wallingford, England. 832 pp.
Boucek, Z. 1988b.
An overview of the higher classification of the Chalcidoidea (Parasitic Hymenoptera). Pages 11-23 in Gupta, V.K. (ed.). Advances in Parasitic Hymenoptera Research. E.J. Brill, Leiden, The Netherlands. 546 pp.
Boucek, Z. and R.R. Askew. 1968.
Index of world Tetracampidae(Hym. Chalcidoidea). Index of Entomomophagous Insects. Delucchi, V. and G. Remaudiere (eds). Le Francois, Paris, France. 254 pp.
Boucek, Z. and J.S. Noyes. 1987.
Rotoitidae, a curious new family of Chalcidoidea (Hymenoptera) from New Zealand. Systematic Entomology 12: 407-412.
Boucek, Z. and J.-Y. Rasplus. 1991.
Illustrated key to west-Palearctic genera of Pteromalidae: Hymenoptera-Chalcidoidea. INRA, Versailles, France. 144 pp.
Breeuwer, J.A. and J.H. Werren. 1993.
Cytoplasmic incompatibility and bacterial density in Nasonia vitripennis. Genetics 135: 565-5747.
Breeuwer, J.A. and J.H. Werren. 1995.
Hybrid breakdown between two haplodiploid species: the role of nuclear and cytoplasmic genes. Evolution 49: 705-717.
Burks, B.D. 1965.
The North American species of Elasmus Westwood (Hymenoptera: Eulophidae). Proceedings of the Biological Society of Washington 78: 201-207.
Burks, B.D. 1971.
A synopsis of the genera of the family Eurytomidae (Hymenoptera: Chalcidoidea). Transactions of the American Entomological Society 97: 1-89.
Dahms, E. and G. Gordh 1997.
A review of the genera of Australian Encyrtidae (Hymenoptera: Chalcidoidea) descrubed from Australia by A.A. Girault with a checklist of included species. Memoirs on Entomology International 9. 518 pp.
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. 1992.
The life history and larval morphology of Aperilampus (Hymenoptera: Chalcidoidea: Philomidinae), with a discussion of the phylogenetic affinities of the Philomidinae. Systematic Entomology 17: 331-339.
Darling, D.C. 1996.
Generic concepts in the Perilampidae (Hymenoptera: Chalcidoidea): An assessment of recently proposed genera. Journal of Hymenoptera Research 5: 100-130.
Darling, D.C. and T.D. Miller 1990.
Life history and larval morphology of Chrysolampus (Hymenoptera: Chalcidoidea: Chrysolampinae) in western North America. Canadian Journal of Zoology 69: 2168-2177.
Delvare, G. and H.-P. Aberlenc. 1989.
Les insectes d'Afrique et d'Amérique tropicale. Clés pour la reconnaissance des familles. PRIFAS, CIRAD-GERDAT, Montpellier, France. 302 pp.
Delvare, G. and Z. Boucek. 1992.
On the New World Chalcididae (Hymenoptera). Memoirs of the American Entomological Institute 53. 466 pp.
De Santis, L. 1967.
Catalogo de los Himenopteros Argentinos de la Serie Parasitica, Incluyendo Bethyloidea. Provincia de Buenos Aires Gobernacion, Comision de Investigacion Cientifica, La Plata. 337 pp.
De Santis, L. 1979.
Catalogo de los Himenopteros Chalcidoideos de America al sur de los Estados Unidos. Comision de Investigaciones Científicas de la Provincia de Buenos Aires, Publicacion Especial, La Plata. 488 pp.
De Santis, L. 1980.
Catalogo de los Himenopteros Brasilenos de la Serie Parasitica Incluyendo Bethyloidea. Editora da Universidade Federal do Parana, Curitiba. 395 pp.
De Santis, L. 1983 (1981).
Catalogo de los Himenopteros Chalcidoideos de America al sur de los Estados Unidos. Primer suplemento. Revista Peruana de Entomologia 24: 1-38.
De Santis, L. 1989.
Catalogo de los Himenopteros Chalcidoideos (Hymenoptera) al sur de los Estados Unidos. Segundo suplemento. Acta Entomologica Chilena 15: 9-90.
De Santis, L. and P. Fidalgo. 1994.
Catalogo de Himenopteros Calcidoideos. Serie de la Academia nacional de Agronomia y Veterinaria 13. 154 pp.
Doutt, R.L. and G. Viggiani. 1968.
The classification of the Trichogrammatidae (Hymenoptera: Chalcidoidea). Proceedings of the California Academy of Sciences 35: 477-586.
Dowton, M., A.D. Austin, N. Dillon, and E. Bartowsky. 1997.
Molecular phylogeny of the apocritan wasps: the Protrotrupomorpha and Evaniomorpha. Systematic Entomology 22: 245-255.
Gauld, I. and B. Bolton (eds). 1988.
The Hymenoptera. Oxford University Press, Oxford, England. xi + 332 pp.
Gibson, G.A.P. 1986a.
Evidence for monophyly and relationships of Chalcidoidea, Mymaridae, and Mymarommatidae (Hymenoptera: Terebrantes). Canadian Entomologist 118: 205-240.
Gibson, G.A.P. 1986b.
Mesothoracic skeleomusculature and mechanics of flight and jumping in Eupelminae (Hymenoptera, Chalcidoidea: Eupelmidae). Canadian Entomologist 118: 691-728.
Gibson, G.A.P. 1989.
Phylogeny and classification of Eupelmidae, with a revision of the world genera of Calosotinae and Metapelmatinae (Hymenoptera: Chalcidoidea). Memoirs of the Entomological Society of Canada 149. 121 pp.
Gibson, G.A.P. 1993.
Superfamilies Mymarommatoidea and Chalcidoidea. Pages 570-655 in Goulet, H. and J.T. Huber (eds). Hymenoptera of the world: An identification guide to families. Canada Communications Group, Ottawa, Canada. 668 pp.
Gibson, G.A.P. 1995.
Parasitic wasps of the subfamily Eupelminae: Classification and revision of the world genera (Hymenoptera: Chalcidoidea: Eupelmidae). Memoirs on Entomology International 5. 421 pp.go to top
Gibson, G.A.P., J.T. Huber, and J.B. Woolley. 1997.
Annotated keys to the genera of Nearctic Chalcidoidea (Hymenoptera). National Research Council of Canada Research Press, Ottawa, Canada. 794 pp.
Graham, M.W.R. de V. 1969.
The Pteromalidae of northwestern Europe (Hymenoptera: Chalcidoidea). Bulletin of the British Museum (Natural History) Entomology, Supplement No. 16. 908 pp.
Gordh, G., E.E. Grissell, and B.D. Burks. 1979.
Superfamily Chalcidoidea. Pages 743-1043 in Krombein, K.V., P.D. Hurd, Jr., D.R. Smith, and B.D. Burks (eds), Catalog of Hymenoptera of America north of Mexico. Vol. 1. Symphyta and Apocrita (Parasitica). Smithsonian Institution Press, Washington, D.C. i-xvi + 1198 pp.
Graham, M.W.R. de V. 1987.
A reclassification of the European Tetrastichinae (Hymenoptera: Eulophidae), with a revision of certain genera. Bulletin of the British Museum (Natural History), Entomology Series 55. 392 pp.
Graham, M.W.R. de V. 1991.
A reclassification of the European Tetrastichinae (Hymenoptera: Eulophidae): revision of the remaining genera. Memoirs of the American Entomological Institute 49. 322 pp.
Greathead, D.J. 1986.
Parasitoids in classical biological control. Pages 289-318 in Waage, J. and D. Greathead (eds). Insect Parasitoids. Academic Press, London. 389 pp.
Grissell, E.E. 1995.
Toryminae (Hymenoptera: Chalcidoidea: Torymidae) a redefinition, generic classification, and annotated world catalog of species. Memoirs of Entomology, Entomology 2. 470 pp.
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A handbook of the families of Nearctic Chalcidoidea (Hymenoptera): Second Edition, Revised. Entomological Society of Washington, Washington, D.C. 87 pp.
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