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Family PTEROMALIDAE

Female of Pteromalidae
The pteromalids represent one of the largest "families" of the Chalcidoidea, consisting of about
2800 world species of morphologically and biologically diverse habits. In our view, this "family"
is the most artificial of the Chalcidoidea. It seems to be defined primarily by what it does not
have in relation to other families, and yet there are always exceptions. Thus, for example, it does
not have the enlarged hindfemora of the Chalcididae (except Chalcedectini), it does not have the
quadrate (rectangular) pronotum of the Eurytomidae (except Spalanginae), it does not have the
exserted ovi-positor of the Torymidae (except for Cea and Roptrocerus), and so on.

As with any large group of taxa, especially a poorly defined one, the approach to classification
differs widely from taxonomist to taxonomist. For example, Riek (1970) and Burks (1979)
considered ormyrids a subfamily of Pteromalidae, whereas Graham (1969) considered them a
distinct family. Riek also considered eucharitids a subfamily of Pteromalidae, but Burks and
Graham considered them a distinct family. The details become extremely difficult to follow and
are too cumbersome for a handbook such as this. What seems useful to us is to present a series of
groupings which seem to represent natural ones (monophyletic lineages). Whether these are
called families (as some have been) or subfamilies or tribes (as others have been) is largely
irrelevant. What is important, is to realize that the groupings seem to be reasonable
(morphologically and biologically). What they actually represent in terms of evolution and
relationship to each other is speculative. The following scheme does not match the most recent
edition of the Hymenoptera Catalog (Burks 1979). We have used the more recent classification
of Boucek (1988) without attempt at explanation. This classification parallels more closely the
one used in the generic keys to Nearctic Pteromalidae by Boucek and Heydon (in Gibson, et al.
1997).

From a practical point of view, identifying Pteromalidae is not easy. In the Nearctic Region the
generic key by Boucek and Heydon (1997, in Gibson, et al.) provides the only starting place.
Boucek alone (1993b) recognized over 40 genera new to the Nearctic Region. Additional keys to
genera and species are listed below in appropriate places. Sometimes a host approach may work
(e.g., a useful paper for identifying pteromalid parasitoids of Nearctic muscoid Diptera is that of
Rueda and Axtell (1985) which includes over 150 SEM photos of the 10 most common
pteromalids associated with these flies).

One of the most useful reference for pteromalid taxonomy in the Northern Hemisphere is
Graham (1969). His work is a monument to the family and provides a basis for future work
throughout the world. He treats over 800 species in about 200 genera for Northwestern Europe
(British Isles and Scandinavia; but all European genera are included). More recently a key to
West-Palearctic genera of pteromalids by Boucek and Rasplus (1991) has improved upon
Graham, especially by the addition of hundreds of illustrations.

The best work for the Australasian area is Boucek (1988) who treats 28 subfamilies, 235 genera,
and lists all of the species for the region. Keys to the 80 genera of India were published by
Farooqui and Subba Rao (in Subba Rao and Hayat 1985). Yoshimoto and Ishii (1965) gave keys
to 11 genera and 17 species of the family from Micronesia. The Neotropical Region is essentially
untouched.

STATISTICS: Number of world species: about 3000 (about 350 Nearctic); number of world
genera: about 550 (about 230 Nearctic).

BIOLOGY AND DISTINGUISHING CHARACTERS: In the subfamily sections which
follow, biology and morphology are discussed by group rather than being given in a single
source for the entire family. Comments given below concerning relative abundance and hosts are
meant to help in confirming an identification based upon the key. The key attempts to cover all
subfamilies of Pteromalidae, but some are so rare they will not likely be collected. Additionally
there are a few genera which are not currently placed to subfamily and these are briefly
mentioned at the end of the following list.

COMMONLY COLLECTED SUBFAMILIES

Spalangiinae: There are about 10 Nearctic species and about 25-50 world species in 2 genera.
The genus Spalangia was revised by Boucek (1963) at the world level.
BIOLOGY: These wasps are associated almost exclusively with flies, especially those breeding
in animal manures, carrion, and decaying plant tissue (Muscidae, Calliphoridae, Scarcophagidae,
Drosophilidae, and Chloropidae). Eggs are laid through the puparial wall, externally on the
dipterous body. The spalangid larva moves about on the host until it finds a suitable point of
attachment where it begins feeding.
CHARACTERS: Black, ant-like wasps, dorso-ventrally compressed with forward projecting
head and antennae attached at sides of mouth. The metasoma is petiolate, the thorax and head are
usually covered with punctures, and the notauli are complete.
Cleonyminae: There are about 15 Nearctic species in 7 genera, and about 170 world species in
40 genera (Boucek 1958). This is a subfamily of diverse and often bizarre species possibly
related to the Eupelmidae. Although females of eupelmids and cleonymids are different
structurally, males are not always easily told apart. This would seem to bring together the
apparently different lineages of the pteromalid and eupelmid-encyrtid lines. Only the tribe
Chalcedectini has been revised in the Nearctic (Grissell 1991).
BIOLOGY: This group is poorly known biologically, but most seem to parasitize wood-boring
beetles and a few parasitize stem or mud nesting Hymenoptera (e.g. Sphecidae, Megachilidae,
Eumenidae).
CHARACTERS: In general, with the eyes diverging ventrally and the pronotum often relatively
long (from anterior to posterior margin, generally as long as wide). Notauli may or may not be
present. Some species have the hindfemur enlarged but without ventral denticles and the tibia
straight, but the Chalcedectini have enlarged hindfemora with ventral denticles and an arched
hindtibia (chalcidid-like). In some species the forefemur may be enlarged with 1 or more ventral
denticles.
Miscogasterinae: There are about 100 Nearctic species in 25 to 30 genera. The number of world
genera and species is unknown. The members of this subfamily are relatively diverse and not
easily defined as evidenced by Graham (1969) who runs them to 12 different couplets in this 54
couplet key to subfamilies of Pteromalidae. According to Boucek (1988) this sufamily may
eventually be merged with Pteromalinae (see next subfamily). Many papers have been published
recently by Heydon (1988a,b; 1989a,b; Heydon and Boucek 1992; Heydon and Grissell 1988,
Heydon and LaBerge 1988) who is attempting to understand the taxonomy of the group.
BIOLOGY: Most members are parasitic on Diptera, especially Agromyzidae, Cecidomyiidae,
Tephritidae, and Anthomyiidae. A few species are reported from Lepidoptera, Coleoptera, and
Hymenoptera (Cynipidae).
CHARACTERS: Distinguishing morphological characters are discussed under the next
subfamily.
Pteromalinae: There are over 250 Nearctic species in about 100 genera. The number of world
species and genera is unknown. As in the Miscogasterinae, Pteromalinae are a diverse and ill-
defined group. Graham (1969) runs this subfamily to 11 couplets of his 54 couplet key, so that
between the Miscogasterinae (with 12 couplets) and Pteromalinae (with 11), nearly one-half the
couplets are devoted to various morphological groupings of the 2 subfamilies.
BIOLOGY: This group is largely reared from holometabolus insects including numerous
Lepidoptera, Coleoptera (such as Chrysomelidae, Scolytidae, Curculionidae), Diptera (such as
Cecidomyiidae, Chamaemyiidae, Syrphidae, Calliphoridae, and Muscidae), and Hymenoptera
(such as Diprionidae, Ichneumonidae, and Braconidae). A number of species are hyperparasitic
on other parasitic Hymenoptera (e.g. Aphelinus and Aphidiidae) through aphids. A few attack
predaceous Diptera (Syrphidae, Chamaemyiidae) that attack Homoptera. One species of
pteromaline, Dibrachys cavus, has been recorded from about 200 hosts in dozens of families
across several orders.
CHARACTERS: In his discussions of Pteromalinae, Graham stated (1969:352) that he could
not attempt to "... give a formal definition of the group." In general (i.e. many exceptions are
present!), Pteromalinae have the notauli incomplete, the clypeus striate and its anterior margin
more or less straight, 1 hindtibial spur, and a non-petiolate abdomen. Miscogasterinae generally
have complete notauli, the clypeus smooth or reticulate and its anterior margin often with small
teeth, 2 hindtibial spurs, and a petiolate abdomen.
UNCOMMONLY COLLECTED SUBFAMILIES

The following Nearctic subfamilies contain less than 50 species in 30 genera. The species are not
often collected. Rarely they may become locally common (see, for example, Eutrichosomatinae),
but this is the exception. It is not always easy to characterize each group succinctly in terms of
morphology. For the sake of taxonomic completeness and general interest we include some
information about each of them.

Eutrichosomatinae: 2 Nearctic species/2 genera. There are 4 world species in 3 genera which
were reviewed at the world level by Boucek (1974b). He reduced the group from family to
subfamily level.
BIOLOGY: The Nearctic species Eutrichosoma mirabilis is the only one with known hosts,
namely Aleutes tenuipes (Leconte) and Smicronyx tychoides (Leconte) (Coleoptera:
Curculionidae) (Boucek 1974b). This species may be extremely abundant in places, and
thousands of specimens may be collected in a short period of time (e.g. we have taken them
sweeping Parthenium infested with Smicronyx).
CHARACTERS: This subfamily is the only one that has the axillae advanced and almost
meeting medially at the hind margins as well as the body covered with flat, scale-like setae.
Colotrechninae: 5 Nearctic species/4 genera. There are about 10 world species in 5 genera.
BIOLOGY: The only hosts known are for the Nearctic species Elachertodomyiaphloeotribi
(Ashmead), a parasite of twig-boring beetles, and for Colotrechnus ignotus Burks reared from
heads of Asteraceae.
CHARACTERS: There are few genera or species, but the group is distinct based upon the
forward projecting axillae, the stigmal vein which is as short as, or shorter than, the stigma itself,
the postmarginal vein which is about as long as the stigmal vein (much as in the torymids), and
the scutellum which has two submedian, parallel longitudinal grooves.
Macromesinae: 1 Nearctic species/1 genus. There are 4 world species in 1 genus, and
Ghesquiere (1963) published a key to them.
BIOLOGY: The species are parasites of Scolytidae and Curculionidae on coniferous trees
(Graham 1969), and we have seen specimens reared from twigs of the evergreen broadleaf
California-Bay (Umbellularia californica). Macromesus americanus Hedqvist is the only known
Nearctic species of this group.
CHARACTERS: Females of this group are the only pteromalids to have the midtarsi 4-
segmented (males are 5-segmented), and both males and females have 2 grooves on either side of
the face below the eyes (i.e. apparently with 2 malar grooves; other pteromalids have 1 or none).
The only known Nearctic macromesine appears to have a triangular projection at the apex of the
scutellum.
Cerocephalinae: 9 Nearctic species/6 genera. There are about 35 world species in 13 genera.
Keys to world genera of this subfamily were given by Gahan (1946) and Hedqvist (1969). Little
has been done with the Nearctic species, but Grissell (1981) provided a key to Cerocephala.
BIOLOGY: All members, except one, parasitize wood-boring beetles and/or possibly their
braconid parasites. The species Choetospila elegans Westwood is a cosmopolitan parasite of
grain and stored-product beetles (e.g. cigarette beetle: Lasioderma serricorne (L.); drugstore
beetle: Stegobium paniceum (L.); granary weevil: Sitophilus granarius (L.)).
CHARACTERS: Members of this subfamily have a globose head with a tooth-like ridge
projecting from between the base of the antennae. The body is generally smooth and often
yellow or brownish in color.
Diparinae: 11 Nearctic species/6 genera. There are about 85 world species in 25 genera. The
Nearctic members of this subfamily were revised by Yoshimoto (1977a).
BIOLOGY: The biology of this group is unknown, but many species have been collected in leaf
litter or duff by the use of Berlese funnels.
CHARACTERS: This group is usually distinguished in females by the strong, dark bristles that
adorn the head and thorax (much as in many Diptera). In males, the antennal "segments" are
usually very long, thin, and with erect setae. Many females are wingless and look considerably
different than the males. Graham (1969) pointed out that in one species of Dipara the males and
females have been put in different subfamilies. Wingless females may easily be mistaken for
proctotrupoids, and one genus (Trimicrops) was originally placed in the Ceraphronidae (Graham
1969).
Ceinae: 6 Nearctic species/2 genera. According to Darling (1991) there are 8 world species in 2
genera. Ceinae was first reported in the New World by Yoshimoto (1977b). Darling and Hanson
(1986) added new species and reported the first record of the genus Cea in the Nearctic. Darling
(1991) revised the world species of Spalangiopelta
BIOLOGY: The known hosts for this group are European records of plant-mining Diptera
(Agromyzidae, Drosophilidae, Graham 1969).
CHARACTERS: In addition to the extremely low antennal insertion mentioned in the key,
ceines may be distinguished by the propodeal spiracles which are situated halfway between the
front and hind margins of the propodeum. It is possible that an uncommon genus or species from
two other subfamilies might key to this group based upon the low antennal placement: one is a
genus of Asaphinae that can be separated by the carinate cheeks and occiput of the head, and the
other is some members of the Eunotinae that may be told by the distinctly concave back of the
head (see figure below). Both groups also have the propodeal spiracle closer to the anterior
margin of the propodeum whereas in Ceinae it is midway between the front and hind margins.
COLLECTING: Darling and Hanson (1986) suggested that because Ceinae are associated with
leaf litter a Berlese funnel is the best method of collection.
Eunotinae: 8 Nearctic species/6 genera. The world numbers are not known. This is a group of
odd-looking species with relatively odd biological habits for a pteromalid.
BIOLOGY: The subfamily is parasitic (or hyperparasitic through encyrtids) on scale insects
(Coccidae: e.g. Saissetia, Ceroplastes, Lecanium, Eriopeltis) and mealybugs (Eriococcidae:
Eriococcus). The European species Eunotus cretaceus Walker is known to be predaceous as a
larva, eating the eggs of Eriopeltis (Graham 1969).
CHARACTERS: The head has a sharply defined occipital edge with the posterior ocelli
touching it, and the first abdominal tergum is at least half as long as the entire abdomen.
Asaphinae: 7 Nearctic species/4 genera. The world numbers are unknown.
BIOLOGY: Members of the genus Asaphes are hyperparasitic on aphidiid wasps in aphids. The
genus Hyperimerus attacks parasites of pseudococcids and is also associated with Neuroptera
(e.g., Chrysopa, Hemerobius), although exact host-parasite relationships are not known (Burks
1979). Bairamlia species in Europe are parasitic on Siphonaptera larvae (Graham 1969), one
attacking fleas in squirrel nests and the other attacking fleas in bird nests. Hosts for the single
Nearctic species of Bairamlia are unknown.
CHARACTERS: Members of this subfamily are most easily recognized by the well-developed
genal (cheek) carinae that flank the sides of the head.
Leptofoeninae: 1 Nearctic species/1 genus. There are 6 world species in 1 genus. This subfamily
is essentially Neotropical with 1 species in Australia and 1 species extending into California and
Arizona from Mexico. It was first reported from the Nearctic by LaSalle and Stage (1985) who
revised the world species.
BIOLOGY: Although biology is unknown for the subfamily, it has been collected on dead or
felled trees, and the long ovipositor is typical of wasps that parasitize wood-boring larvae
(LaSalle and Stage 1985).
CHARACTERS: These are extremely unusual chalcidoids and could easily be mistaken for
ichneumonids or braconids. The extra-Nearctic species are long (up to almost 4 cm including the
well exserted ovipositor), but the Nearctic one is only 1 to 2 cm. All species are elongate and
thin. In all species the head has well-developed ridges and crests in the area between the eye and
antennal basin.
Herbertinae: 1 Nearctic species/1 genus. World numbers are unknown. Boucek (1988)
proposed this subfamily for the genus Herbertia which occurs throughout the world.
BIOLOGY: These small wasps are parasites of leaf-mining Diptera (Boucek 1988).
CHARACTERS: This subfamily is recognized by the dense setae that cover the head (including
eyes), the dorsum of the mesosoma, and the wings. In addition, the first gastral tergum is half the
length (or more) of the abdomen, and is polished dorsally.
Unplaced taxa: In addition to the above subfamilies there remain a few odd genera that are not
currently placed to subfamily. These are relatively unlikely to be collected except possibly for
Hemadas nubilipennis, a phytophagous, gall-forming species on blueberries in eastern North
America. It may be recognized by a darkened spot on the fore-wing and the scutellum that
projects over the propodeum and obscures it in dorsal view.

A few other subfamilies are known, each of which contain a few genera or less with one species
each. These include Erotolepsiinae, Ormocerinae, Pireninae, and Panstenoninae. Panstenoninae
was listed by Burks (1979) as occurring in the Nearctic Region based upon 1 specimen. Boucek
(1988) removed that species to another subfamily but replaced it with a widespread species,
Panstenon poaphilum, which was described in Heydon and Boucek (1992).

COLLECTING: Pteromalids, being a group rich in numbers and diversity, may be collected
nearly anywhere. All of the sorts of localities mentioned for other chalcidoids will be found to
harbor pteromalids. Sweeping meadows, shrubs, trees, pondside vegetation, and individual
patches of herbaceous and annual plants (especially in flower) will yield pteromalids. Holding
batches of composite seed heads, seed pods of legumes, dead twigs, cones, galls, lepidopterous
cocoons, dipterous puparia from manure or carcasses, mummified aphids, nests of aculeate bees
and wasps, will all produce pteromalid specimens.
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Last Updated:01/08/2003 16:16:08