family pteromalidae

7/27/2019 Family Pteromalidae

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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 forCea andRoptrocerus), 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 toeach 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)


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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 forNorthwestern 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 onthe host until it finds a suitable point of attachment where it begins feeding.

CHARACTERS: Black, ant-like wasps, dorso-ventrally compressed with forwardprojecting 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-


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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 toBoucek (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). Anumber 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 apetiolate abdomen.


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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 taxonomiccompleteness 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 speciesEutrichosoma mirabilis is the only one with known

hosts, namelyAleutes 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 sweepingParthenium 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 Hedqvistis 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.


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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, theantennal "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 ofDipara 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 ofSpalangiopelta

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 thekey, 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 whereasin 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 speciesEunotus cretaceus Walker is known

to be predaceous as a larva, eating the eggs ofEriopeltis (Graham 1969).


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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 genusAsaphes are hyperparasitic on aphidiid wasps inaphids. The genusHyperimerus 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 ofBairamlia 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. Thissubfamily 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 genusHerbertia 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 forHemadas 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


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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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family pteromalidae

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