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 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
Literature on chalcid wasps is vast. Probably the two most important
publications for any serious student of chalcidology in North America are:
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significantly outdated but still essential section on Chalcidoidea in Catalog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.
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
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
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).
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,
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
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
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.
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:
- an externally visible prepectus
- mesothoracic spiracle at exposed lateral edge of mesoscutum
- flagellum with uniquely structured longitudinal (multiporous
plate) sensilla [SEM]
He also proposed that Chalcidoidea was the sister group of the family
Mymarommatidae based on three other features shared by the two taxa:
- unique loss of two of the three ancestral (plesiomorphic) sites
of origin for a mesothoracic muscle, the mesotrochanteral depressor
- origin of an anteriorly projecting
phragma as the site of origin
for all or part of the remaining portion of the mesotrochanteral muscle,
the mesotergal-mesotrochanteral (t2-tr2) muscle, and
- loss of the basal ring from the male
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
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
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
[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
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.
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
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
- 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
- Gibson (1986a)
- Woolley (1988)
- Grissell (1995)
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),
- Delvare and Boucek (1992),
Wijesekara (1997), Narendran
- 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
- Heraty (1985), Heraty
- Graham (1987), Graham
- Gibson (1989), Gibson
- Burks (1971), Stage
and Snelling (1986), Zerova (1988),
- Boucek (1974)
- Annecke and Doutt (1961),
Subba Rao and Hayat (1983),
Schauff (1984), Huber
(1986), Viggiani (1988),
Noyes and Valentine (1989a),
- Boucek (1986), Hanson
- Boucek (1978), Darling
- Graham (1969), Boucek
and Rasplus (1991)
- Boucek and Noyes (1987)
- Woolley (1986), Woolley
- LaSalle (1987)
- Boucek and Askew (1968),
- Narendran (1994), Grissell
- Doutt and Viggiani (1968),
Yousuf and Shafee (1986a),
Yousuf and Shafee (1986b),
Yousuf and Shafee (1987)
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.
and Schauff (1997a) in * Gibson et
al. (1997) - Nearctic region
and Schauff (1997b) - Nearctic region
and LaSalle (1995) - Costa Rica
(1991) - Australia
and Valentine (1989b) - New Zealand (plus keys to genera of 12 of 16
and Aberlenc (1989) - tropical Africa and America
(1988a) - Australasian region (plus keys to genera of 14 of 21
and Bolton (1988) - Britain
Rao and Hayat (1985) - Oriental region
(1984) - Canada
(1980) - Ethiopian region
(1978) - European USSR
and Hernandez (1978) - Cuba
(1970) - Australia
Boucek, and Hoffer (1964) - Europe
(1969) - Europe (plus key to genera of Pteromalidae)
(1952) - European USSR
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
- Central and South America
- De Santis (1967), De
Santis (1979), De Santis (1980),
De Santis (1983), De
Santis (1989), De Santis and Fidalgo
- Alayo (1970)
- Subba Rao and Hayat (1986)
- Baltazar (1966)
- Alayo D. Pastor 1970.
- Catalogo del los Himenopteros de Cuba. Editorial Pueblo y
Educacion. 218 pp.
- Alayo D. Pastor and L.R. Hernandez.
- 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.
- Morphology of the antenna cleaner in the Hymenoptera with particular
reference to non-aculeate families (Insecta). Zoologica Scripta
- Bendel-Janssen, M. 1977.
- Zur Biologie Okologie und Ethlogie der Chalcidoidea. Mitteilungen
aus der Biologischen Bundesanstalt für Land- und Forstwirtshaft:
- 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
- Burks, B.D. 1971.
- A synopsis of the genera of the family Eurytomidae (Hymenoptera:
Chalcidoidea). Transactions of the American Entomological Society
- 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.
- 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
- 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
- De Santis, L. 1989.
- Catalogo de los Himenopteros Chalcidoideos (Hymenoptera) al sur de
los Estados Unidos. Segundo suplemento. Acta Entomologica Chilena
- 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
- 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
- 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.
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updated June 18, 1998 by J.D. Read, BRP/ECORC
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