About Chalcidoidea (Chalcid wasps)
Chalcidoid or chalcid wasps are
one of the most diverse groups of Hymenoptera (bees, ants, wasps) numerically,
structurally, and biologically. They range in size from the smallest insect
known, Dicopomorpha echmepterygis Mockford (1997), at about
130 microns (0.13 mm), to over 30 mm, including bizarre as well as beautiful
winged and wingless forms. About 22,000 valid species have been described in
about 2,100 genera world-wide, but 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. There are over 2,600 described species in over 700 genera
in North America. Illustrated keys to the families and genera of chalcids known
from North America are given in Annotated
Keys to the Genera of Nearctic Chalcidoidea (Hymenoptera) by Gibson et al. (1997).
Other important references to the chalcid taxonomic literature, with emphasis on
the North American fauna, are given in the sections below.
Literature on chalcid wasps is vast, but there are two vital resources for
any serious student of chalcidology, biological control specialist, or anyone
else who needs information on the literature, names, distribution or hosts of
world chalcids. The Universal Chalcidoidea
Database by John Noyes provides an online version of his earlier CD-Rom
version of the database (Noyes
2002). Although the CD-Rom and Internet databases are not official
publications under the International Code of Zoological Nomenclature, they are
unique and invaluable. They database about 31,000 world chalcid species names as
well as over 550,000 entries to some aspect of the taxonomy, biology, morphology
and distribution of all valid species. Each data entry is validated by being
linked to one or more of over 40,000 original chalcid literature references
published from 1758 through 2003. Both the CD and Internet versions also contain
over 350 chalcid
images of species. The Internet version additionally contains an online
illustrated key to the 19 recognized world chalcid families and family
information, including diagnostic features, subfamily classification,
biological attributes and major references. Another extensive source of online
information concerning the systematics and biology of chalcids can be found on
subsite of the Systematic Entomology
Laboratory web site. The Chalcidoidea section consists of an online version
of Grissell and Schauff
(1997b) (excluding the pictorial key to families) and PDF documents of all
24 issues of Chalcid
Forum, an international newsletter dedicated to promoting communication
among chalcid workers. The SEL site also contains information on equipment and
methods used to collect, rear and preserve chalcids for study under Collecting
Chalcidoidea: Why and How. Because of their small size and often extreme
fragility, special techniques are required to preserve specimens for microscopy.
A detailed, step by step procedure for mounting minute chalcids on microscope
slides is online as part of Collecting
and Preserving Chalcidoids. Another site with instructions for preparing
slides of chalcids is Encarsia
of Australia. Comprehensive information on the morphology of chalcids and
the terms used by taxonomists to describe chalcid species is found in Chalcid Wasps:
Introduction to Glossary of Positional and Morphological Terms. A final
online resource for information about chalcids is Nomina Nearctica,
which provides a checklist of valid names of described species of Chalcidoidea
and other Hymenoptera in America north of Mexico based on alphabetical lists of
families, genera and species.
Within the United States, training in systematics is available at several
universities. The Departments of Entomology at Texas A&M University in
College Station and at the University of Riverside, California, currently have
several students studying chalcids and other groups of parasitic Hymenoptera.
Information on the people and programs can be found under the Parasitic Hymenoptera Research
Laboratory at Texas A&M and under Research on Chalcidoid
Systematics at the University of Riverside. A directory of world
chalcidologists is available at http://iris.biosci.ohio-state.edu/newsletters/cmen.html.
Economic Importance and Biological
DiversityAlthough often very beautiful because of their unusual structures
and striking colors or metallic hues, chalcids are very tiny and unobtrusive,
not biting or stinging humans, living in their domiciles, or carting off their
food. Because of this they remain largely unnoticed. They are unknown to all but
the specialist and are under-appreciated for their role in the environment and
benefit to man [see LaSalle
and Gauld (1992) and LaSalle (1993)]. About 80
chalcid species are known to be pests of agriculture (mostly seed-feeders in the
families Eurytomidae and Torymidae) and some chalcids are considered harmful
because they are hyperparasitoids, but most are economically and environmentally
beneficial. The large majority of chalcid species are primary parasitoids of
other insects and arachnids and as such they are important participants in
nature's own control system for regulating arthropod populations. In addition to
the largely unappreciated role of most species in helping to control what might
otherwise be pest species, over 800 chalcid species have been associated with
targeted biological control programs. This represents about two-thirds of all
biocontrol programs involving Hymenoptera, and about one-third of all biocontrol
programs in which partial or complete economic control of an insect pest was
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 in about 340 families of 15 insect orders (Blattaria, Coleoptera,
Diptera, Hemiptera, Homoptera, Hymenoptera, Lepidoptera, Mantodea, 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.
Agaoninae (Agaonidae) are exclusively phytophagous within the ovarioles of figs
(Ficus), but are beneficial as the obligate pollinators of figs. Some
members of five other families (Eulophidae, Eurytomidae, Pteromalidae,
Tanaostigmatidae and Torymidae) are seed-feeders or gall-formers on plants,
though for many chalcids reared from galls it is not known whether they are
primary gall-formers, or inquilines or parasitoids in the galls. John Noyes
provides a concise summary of known chalcid biology by family in Chalcid Forum 24.
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 parasitoids,
as primary or hyperparasitoids, 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
IdentificationBecause of their small size
and structural diversity it is difficult to give easily visible and reliable
features that differentiate 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. This vein
usually branches apically [figure]
so that the vein complex typically consists of a submarginal,
vein. Unfortunately, some other parasitic microhymenoptera, most notably the
family Scelionidae (Platygastroidea), have a forewing venation that is very
similar to typical chalcids and consequently they are often mistaken for
chalcids. Many chalcids are quite easily distinguished by having a distinct
metallic colouration, which almost all other microhymenoptera with a similar
venation lack. However, many chalcids, particularly apterous and brachypterous
ones, are non-metallic and somewhat more arcane features are needed to
differentiate these from other microhymenoptera. Most chalcids have the pronotum
more or less distinctly separated from the base of the forewing because there is
an intervening sclerite, the prepectus,
between the pronotum and mesopleuron
Although often difficult to see, chalcids also have a unique placement of the mesothoracic
spiracle — it is located between the pronotum and the lateral margin of the
often above the anterodorsal angle of the prepectus but at least separate from
the anterodorsal margin of the mesopleuron (mesepisternum)
Other microhymenoptera 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. Furthermore, other
microhymenoptera with a forewing venation similar to chalcids have the
posterolateral angle of the pronotum
extending back to touch the more or less oval sclerite (tegula)
that covers the base of the forewing. As a result of this structural difference,
most chalcids have the pronotum moveable relative to the mesothorax, whereas the
pronotum is rigidly associated with the mesothorax in other microhymenoptera.
Finally, almost all chalcids also have ridge-like, longitudinal
sensilla on the flagellum of the antenna [SEM].
Superfamily RelationshipsGibson (1986a) proposed
that Chalcidoidea, including the family Mymaridae, is monophyletic based on
three shared derived features (synapomorphies) that are listed above as
and Huber (2000) subsequently discovered that individuals of the family
Rotoitidae have a linear prepectus that normally is concealed under the
posterolateral margin of the pronotum. This is the hypothesized groundplan
(primitive) structure for Apocrita (Gibson 1999) and would
invalidate the first hypothesis if the rotoitid-like structure is the groundplan
state for the superfamily and not an autapomorphic (unique) secondary reduction
of Rotoitidae alone.
- an externally visible prepectus [SEM]
- mesothoracic spiracle at exposed lateral margin of mesoscutum [SEM],
- flagellum with uniquely structured longitudinal (multiporous plate)
Gibson (1986a) also
proposed that Chalcidoidea was the sister group of the family Mymarommatidae
based on three other features shared by the two taxa:
Since then most authors have classified mymarommatids
separately as their own superfamily, the Mymarommatoidea.
- 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 genital
Quicke et al.
(1994) also provided evidence that chalcids other than Mymaridae (and
Mymarommatidae) 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].
Furthermore, they have a transversely striated band of notal
membrane, the laminated
bridge near the base of the second valvulae. Quicke et al. proposed that
both the unique structure of the second valvulae and the laminated bridge were
synapomorphies uniting all chalcids except mymarids. This would support an
hypothesis that Mymaridae is the sister group of all other Chalcidoidea. Gibson and Huber (2000)
subsequently showed that females of at least one of two known genera of
Rotoitidae have an ovipositor structure that is intermediate between the
mymarid-like and other chalcid-like structures. The rotoitid ovipositor has
about the basal half of the second valvulae structured like mymarids and most
other Hymenoptera, but the apical half with asymmetrical overlapping portions
similar to other chalcids. This intermediate structure could be evidence that
Rotoitidae is the second-most basal lineage of Chalcidoidea after Mymaridae.
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, chalcids have this muscle attached along the
entire length of the laterophragmal apodeme and this is additional evidence for
excluding Mymarommatidae from Chalcidoidea. Heraty et al. 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
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
are uniquely structured in Chalcidoidea. The setae were said to be "distinctly
flattened" in chalcids and cylindrical in the rest of the Hymenoptera, though
the state in mymarommatids was not stated explicitly. These features need to be
examined more comprehensively throughout 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, relationships between Mymarommatidae +
Chalcidoidea and other apocritan Hymenoptera are more controversial. Rasnitsyn (1988) proposed
that Chalcidoidea is the sister group of the Platygastroidea (Platygastridae +
et al. (1999) reanalysed Rasnitsyn's 1988 data using cladistic parsimony
analysis and, perhaps not surprisingly, retrieved a (Chalcidoidea +
Mymarommatidae) + (Platygastridae + Scelionidae) sister-group relationship from
most analyses. Much more unexpected was that Ceraphronoidea was also indicated
to be closely related to Chalcidoidea + Mymarommatidae, either as the sister
group of the clade (Chalcidoidea + Mymarommatidae) + (Platygastridae +
Scelionidae) or as the sister group of Platygastridae + Scelionidae. They
suggested that these indicated relationships might reflect parallel reductions
correlated with small body size, particularly in the large number of wing
venational characters used in the analysis.
Comparatively recently within systematics, molecular evidence has been used
to supplement more traditional morphological studies for analysing phylogenetic
relationships. A Chalcidoidea + Platygastroidea sister-group relationship was
supported through analysis of the mitochondrial 16S rRNA gene by Dowton et al. (1997),
and this was also a commonly retrieved relationship in the much more
comprehensive study of Dowton and Austin (2001).
They analysed three gene regions for many more apocritan taxa, plus morphology
based primarily on Ronquist et al. (1999).
Maximum parsimony analysis of just the molecular data with all character
transformations weighted equally indicated Chalcidoidea as the sister group of
Monomachidae + (Diapriidae + Maamingidae), with this group part of an unresolved
trichotomy comprised of Proctotrupidae + Vanhorniidae and Cynipoidea +
Platygastroidea. Simultaneous analysis of the molecular and unordered
morphological data resulted in a Cynipoidea + Chalcidoidea sister-group
relationship, which formed an unresolved polychotomy with most other apocritan
groups. However, almost all other analyses of the molecular evidence alone or
simultaneous analysis of the molecular evidence and morphology retrieved a
Chalcidoidea + Platygastroidea sister-group relationship, which inevitably
comprised the terminal clade within Apocrita. Relationships of this clade with
other apocritans varied considerably within the different analyses and Dowton and Austin (2001)
should be examined for specifics. Based simply on morphology, Gibson (1999) proposed that
Platygastroidea forms a monophyletic group with Pelecinidae + Proctotrupidae +
Vanhorniidae, and that Chalcidoidea + Mymarommatoidea likely represent a
relatively basal clade with uncertain relationships in Apocrita.
Family RelationshipsIf anything, higher
level relationships within Chalcidoidea are even less well resolved than are
relationships of Chalcidoidea with other Hymenoptera. A comprehensive historical
review of chalcid higher classification was given by Boucek (1988b).
Historically, family classification has been based primarily on external
morphology rather than phylogenetic relationships. Because of this, and because
of the extreme structural diversity that characterizes the group, there have
been a comparatively large number of families recognized within the superfamily
as well as instability in the number of families recognized. Anywhere from 9 to
24 families have been recognized since about 1950, with 19 or 20 families
generally being recognized at present. However, monophyly of many if not most of
the families is in doubt. Chalcid families 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. As one might suspect,
the classification of some subfamilies to one family or another is controversial
and there is even uncertainty about the proper family classification of some
Although still at what might be called an embryonic stage, molecular analyses
have now started to provide independent evidence to test and refine family
concepts that are widely acknowledged to be less than satisfactory. Rasplus et al. (1998)
concluded from analysis of the D1 and D2 regions of the 28S rRNA gene that
Agaonidae sensu Boucek (1988a) is not
monophyletic. They redefined the family to include only the pollinating
Agaoninae sensu Boucek (1988a), reassigning some subfamilies to the
Pteromalidae and excluding other subfamilies that they left unassigned to
family. More recently, Gauthier et al. (2000)
analysed the D2 region of the 28S rDNA gene and concluded that the family
Elasmidae deserved only tribal status within the subfamily Eulophinae of
Eulophidae. As for the previous study, they also excluded some taxa from
Eulophidae and left them unassigned to family. Campbell et al.
(2000) provided the first comprehensive analysis of chalcid subfamily and
family relationships based on analysis of the D2 region of 28S rDNA. They
included 85 taxa, representing 18 families and 32 subfamilies, and concluded
that their analysis placed 80% of the taxa (including outgroup taxa) "into some
form of realistic grouping (generic or family group taxon) based on
morphological evidence". Of 12 families with more than one taxon represented,
only three were indicated as monophyletic (Eucharitidae, Mymaridae and
Trichogrammatidae), though Eulophidae was monophyletic with the inclusion of
Elasmus (= Elasmidae). More unrealistic or intriguing results included
none of the three genera of Eunotinae (Pteromalidae) grouping together, and
neither Eupelminae nor Calosotine (Eupelmidae) being indicated as monophyletic
or showing any affinities with Cleonyminae, Tanaostigmatidae or Encyrtidae.
Interestingly, their analysis indicated Mymaridae as the sister group of all
other Chalcidoidea, whereas most analyses of Dowton and Austin (2001)
retrieved Mymaridae as an apical clade within Chalcidoidea. Resolving the
relationships of Mymaridae with other Chalcidoidea is critical for establishing
the ancestral life history of the group. Traditionally, chalcids have been
considered as most likely evolving from some ectoparasitoid of wood-boring
beetles. If Mymarommatoidea is the sister group of Chalcidoidea and if Mymaridae
is truly the sister group of all other Chalcidoidea then parsimony would
indicate the immediate ancestor was an endoparasitic egg parasitoid Dowton and Austin
There is no doubt that molecular analysis of chalcid relationships is an
exciting new frontier that holds promising rewards for our understanding of the
evolution of the group. However, such analyses seem likely to add to the
instability of chalcid classification, at least in the short term, and by
themselves seem unlikely to provide a fully resolved pattern of chalcid
relationships. In order to fully resolve the evolutionary history of the
Chalcidoidea it will also be necessary to develop a comprehensive, accurate,
morphological-based character matrix for the superfamily. This is an extremely
complex and daunting task considering the enormous diversity of the group, but
must be done in order to advance chalcid classification beyond the level of
personal preference. Although no cladistic hypotheses of family relationships
based on analysis of morphology have considered the Chalcidoidea in its
entirety, Noyes (1990)
presented a tree diagram illustrating one set of potential chalcid family
relationships (see reproduction in "A word on chalcidoid
classification" or dendrogram 1 in Heraty et al. 1997). Gibson et al. (1999)
reviewed current concepts of chalcid phylogenetics and classification, and this
paper should be consulted for a comprehensive list of relevant publications.
However, some of the more major molecular or morphological analyses that have
investigated monophyly and relationships of chalcid families or subfamilies
through explicit character state analysis are listed below. References are given
under the primary family or families investigated though the papers often
include discussion of other families.
- Rasplus et al.
- Shaffee and Rizvi
(1990), Heraty et
al. (1997), LaSalle
et al. (1997)
- Chalcididae and/or Leucospidae
- Delavare 1988, Wijesekara 1997, Delvare 1999
- Encyrtidae, Eupelmidae, and/or Tanaostigmatidae
- LaSalle and Noyes
- Eucharitidae and/or Perilampidae
- Heraty and Darling
(1989), Darling and
Miller (1990), Darling
- Eulophidae and Elasmidae
- Gauthier et al.
- Schauff (1984), Gibson (1986a)
- Dzhanokmen (2000),
Torok and Abraham
- Woolley (1988)
Recognized Families and Major Taxonomic
ResourcesListed below in alphabetical order are the 19 families of
Chalcidoidea that are generally accepted. Five of the family names are linked to
other Internet pages developed by specialists that provide some type of
information on the respective families. 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)
- Yasnosh (1976), Hayat and Verma (1980),
Hayat (1983), Yasnosh (1983), Hayat (1994), Hayat (1998)
- Narendran (1989),
Delvare and Boucek
- Tachikawa (1963),
Trjapitzin (1973a), Trjapitzin (1973b), Trjapitzin and Gordh
and Gordh (1978b), Prinsloo and Annecke
(1981), Noyes and
Hayat (1984), Noyes
(1988), Dahms and
Gordh (1997), Noyes
- Heraty (1985), Heraty (1994), Heraty (2002)
- Eulophidae (including Elasmidae)
- Burks (1965), Graham (1987), Graham (1991), Hansson (2002)
- Gibson (1989), Gibson (1995)
- Burks (1971), Stage and Snelling
- Boucek (1974)
- Annecke and Doutt
(1961), Subba Rao and
Hayat (1983), Schauff
(1988), Noyes and
Valentine (1989a), Yoshimoto (1990)
- Boucek (1986), Hanson (1992), Narendran (1999)
- Boucek (1978), Darling 1996
- Graham (1969), Boucek and Rasplus
- Boucek and Noyes
(1987), Gibson and
- Woolley (1986), Woolley (1988)
- LaSalle (1987)
- Boucek and Askew
- Narendran (1994),
Grissell (1995), Grissell (1999)
- Doutt and Viggiani
(1968), Yousuf and
Shafee (1986a), Yousuf and Shafee
(1986b), Yousuf and
Shafee (1987), Pinto
Regional Keys to FamiliesIt 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. Gibson (1993)
provided a world key that attempted to include exceptional forms, but because of
this the key is quite complicated and it is recommended that individuals
normally try to identify families using relevant regional keys. An online key to
the families of Chalcidoidea and Mymarommatoidea is given by Noyes
(2003). 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 alphabetical order by area covered; those with an asterisk also
include keys to genera.
Australasian region — * Boucek (1988a) (plus keys
to genera of 14 of 21 families)
Australia — Riek (1970); Naumann (1991)
Gauld and Bolton
Canada — Yoshimoto (1984)
Rica — * Hanson and LaSalle
Cuba — * Alayo
and Hernandez (1978)
Ethiopian region — Prinsloo (1980)
* Peck, Boucek, and Hoffer
(1964); Graham (1969)
(plus key to genera of Pteromalidae)
European USSR — * Nikolskaya (1952); * Medvedev (1978)
region — Grissell and
Schauff (1997a) in *
Gibson et al. (1997); Grissell and Schauff
New Zealand — * Noyes and Valentine
(1989b) (plus keys to genera of 12 of 16 families)
Oriental region — * Subba Rao and Hayat
Tropical Africa and America — Delvare and Aberlenc
World — Gibson
CatalogsCatalogs 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. The CD-Rom catalog
of Noyes (2002) and the
online catalog of Noyes
(2003) basically supersede previously published printed catalogs. However,
listed below are some of the more recent published catalogs for Chalcidoidea;
catalogs for individual families or genera are not listed.
Canada and United States — Peck (1963), Gordh et al.
Central and South America — De Santis (1967), De Santis (1979), De Santis (1980), De Santis (1983), De Santis (1989), De Santis and Fidalgo
Cuba — Alayo
India — Subba
Rao and Hayat (1986)
Philippines — Baltazar (1966)
- Alayo D. Pastor. 1970.
- Catalogo del los Himenopteros de Cuba. Editorial Pueblo y Educacion. 218
- 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.
- Baltazar, C.R. 1966.
- A catalogue of Philippine Hymenoptera (with a bibliography, 1758-1963).
Pacific Insects Monograph: 8: 1-488.
- Basibuyuk, H.H. and D.L.J. Quicke. 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), Supplement23.
- 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 Scandinavica9:
- 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.
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