The Pireninae

The chalcidoid wasps are truly a remarkable array: tiny wonders coming in a bewildering variety of forms. For this post, I'm looking at the members of the chalcidoid subfamily Pireninae.

Macroglenes sp., copyright Charley Eiseman.

The Pireninae are currently recognised as one of the subfamilies of the Pteromalidae, a chalcidoid 'family' that is long overdue for reclassification as phylogenetic studies have agreed that it is extensively polyphyletic* (e.g. Heraty et al. 2013). The pirenines are very small wasps, about one to two millimetres in length. They've always struck me as having a fairly fly-like habitus: they lack the metallic coloration and strong sculpturing of many other pteromalids, often being uniformly black or yellow, and carry upright bristle-like setae on the head and mesosoma. Characteristic features of the Pireninae also include antennae inserted low on the face, reduced numbers of antennal segments (and hence often rather short antennae), a large clypeus that often protrudes ventrally, and a dorsally rounded mesosoma often with deeply impressed notauli (longitudinal grooves on the mesoscutum) (Bouček 1988). About ten genera are currently recognised in the subfamily. Perhaps the most remarkable is the genus Zebe, named by John La Salle in 2005 from a single female that he says, at the time, had stymied multiple hymenopterists as to what it might be for two decades. Zebe has legs with four-segmented tarsi, instead of the five-segmented tarsi of other pteromalids, and the female has a long horn extending forward from the mesoscutum and hanging over the head. As in other micro-wasps with comparable structures, this horn probably provides space for the retraction of an extraordinarily long ovipositor.

*But just in case anyone who stands to have some influence is reading, I will point out that the number of subfamilies that need to excised from the Pteromalidae could be kept to a minimum if the family is expanded to include the Ormyridae, Torymidae, Eucharitidae and Perilampidae. Bonus points that these are the families that are the hardest to distinguish from pteromalids to begin with.

Female Zebe cornutus, from Mitroiu (2011).

The life habits of most pirenines are very little known. Those whose hosts are known develop as parasites of Cecidomyiidae, gall midges, and may be found in association with the galls produced by those flies on various plants. Species of the genus Macrogelenes attack cecidomyiids associated with grasses, and some have been investigated as control agents for midges on commercial crops. So these tiny little wasps could prove themselves very valuable to humans.

REFERENCES

Bouček, Z. 1988. Australasian Chalcidoidea (Hymenoptera): A biosystematic revision of genera of fourteen families, with a reclassification of species. CAB International: Wallingford (UK).

Heraty, J. M., R. A. Burks, A. Cruaud, G. A. P. Gibson, J. Liljeblad, J. Munro, J.-Y. Rasplus, G. Delvare, P. Janšta, A. Gumovsky, J. Huber, J. B. Woolley, L. Krogmann, S. Heydon, A. Polaszek, S. Schmidt, D. C. Darling, M. W. Gates, J. Mottern, E. Murray, A. D. Molin, S. Triapitsyn, H. Baur, J. D. Pinto, S. van Noort, J. George & M. Yoder. 2013. A phylogenetic analysis of the megadiverse Chalcidoidea (Hymenoptera). Cladistics 29: 466–542.

La Salle, J. 2005. Zebe cornutus gen. et sp. nov., a new Pireninae (Hymenoptera: Pteromalidae) with 4-segmented tarsi and a mesoscutal horn. Acta Societatis Zoologicae Bohemoslovenicae 69: 193–197.

The Life and Times of Diaulomorpha

Diaulomorpha is a fairly typical genus of the diverse micro-wasp family Eulophidae. Like most other eulophids, members of this genus are slender with a relatively soft metasoma. The mesosoma, on the other hand, is tougher, weakly vaulted, and conspicuously reticulate dorsally. Members of the genus are known from Australasia and South America (Bouček 1988).

Body of female Diaulomorpha itea, from Bouček (1988).

Diaulomorpha species are parasitoids of insect larvae that live as miners in leaves. They are known to feed on both Lepidoptera and Hymenoptera larvae; it seems that it is not the identity of the host that attracts them but the lifestyle. They are multivoltine, that is they can go through multiple generations in the course of a year. The breeding cycle and behaviour of a Diaulomorpha species was described by Mazanec (1990) as a parasitoid of the jarrah leafminer Perthida glyphopa, a moth whose larvae attack the leaves of jarrah Eucalyptus marginata.

Mating between males and females occured after a brief courtship ritual in which the pair each extended their wings upwards and beat them up and down. Females located host larvae by running across the leaf surface and drumming the outside of prospective mines with their abdomen. They would then drill into the mine with their ovipositor, though of course the host larva would generally be trying to escape the wasp's attentions; a female might have to drill several holes before successfully piercing the caterpillar. The ovipositor would then be 'stirred' into the host to cause haemolymph and other fluids to leak out of its skin, and the wasp would feed on this fluid through the hole formed by the ovipositor. Egg laying would begin shortly after the wasp had finished feeding. Egg production was relatively slow, with only five or six eggs able to develop within the mother at a time, and the female wasp would lay through a newly created hole into the mine near the selected host. Usually only one egg would be laid in a mine but sometimes multiple eggs would be laid and the emerging larvae would share the host individual. After laying an egg, the female would tap around the laying hole with the tip of her metasoma, presumably depositing some chemical that would signal to other Diaulomorpha females that the mine had already been attacked. The host, meanwhile, would stop feeding after being stabbed with the female's ovipositor and would finally die after about a day and a half. It was around this time that the larva(e) would hatch and commence feeding on its remains.

Though healthy hosts would obviously be preferable, female Diaulomorpha were not above attacking hosts that had already died or had already been parasitised by other wasps. Such hosts were particularly likely to be attacked by young females that had not yet learnt to deal with the defensive actions of a healthy host. Deceased hosts could present a problem in that their bodily fluid pressure had been lost, and the female might have to stab them with her ovipositor several times before she had ingested enough fluid to begin laying. Pre-parasitised hosts were less of a problem as endoparasitic wasp larvae within the host would die after the Diaulomorpha's stab along with the host.

Mazarec (1990) found parasitism levels by Diaulomorpha within the host population to be low. What is more, as host populations increased the level of parasitism would plateau, so the proportion of parasitised hosts was far lower in dense host populations. This presumably resulted from the wasp's low rate of egg production: as host populations increased, the population of wasps did not keep up with it. As such, the role of Diaulomorpha in pest control is probably limited.

REFERENCES

Bouček, Z. 1988. Australasian Chalcidoidea (Hymenoptera): A biosystematic revision of genera of fourteen families, with a reclassification of species. CAB International: Wallingford (UK).

Mazanec, Z. 1990. The immature stages and life history of Diaulomorpha sp. (Hymenoptera: Eulophidae), a parasitoid of Perthida glyphopa Common (Lepidoptera: Incurvariidae). Journal of the Australian Entomological Society 29: 147–159.

Perilampella acaciaediscoloris: An Australian Gall Wasp

The small but incredibly diverse chalcidoid wasps are mostly known as parasitoids, their larvae attacking the eggs and young of others insects. Some, however, have chosen the vegetarian option, inserting their eggs into plant rather than animal tissue. As the larva develops, it induces the host plant to develop an often bizarre-looking growth around it that provides both shelter and food; this growth is known as a gall.

Antenna, forewing venation and dorsum of Perilampella acaciaediscoloris, from Bouček (1988).

Perilampella acaciaediscoloris is an gall-forming wasp that was first described by Froggatt in 1892 from galls that he collected on the wattle Acacia discolor, a species now regarded as a synonym of the sunshine wattle A. terminalis of south-eastern Australia. Froggatt placed his species in the genus Cynips (which is not part of the Chalcidoidea but belongs to a different micro-wasp superfamily, the Cynipoidea) but it is now placed in the chalcid subfamily Ormocerinae in the (polyphyletic) Pteromalidae. Ormocerinae are fairly generalised 'pteromalids' that are non-metallic in colour and often finely sculpted. So far as is known, ormocerines are all associated with galls in one way or another, either as gall-causers themselves or as inquilines (species that lay their eggs in the galls caused by other insects). Another ormocerine species, Trichilogaster acaciaelongifoliae, has been introduced from Australia to South Africa to help control the Sydney golden wattle Acacia longifolia.

The related ormocerine Trichilogaster acaciaelongifoliae, copyright Simon van Noort. Perilampella acaciaediscoloris most obviously differs from this species in its hairier and darker wings, and more shiny mesosoma.

The genus Perilampella differs from other ormocerines in being particularly shiny, with little clear setation. Bouček (1988) listed four Australian species in the genus, noting that P. acaciaediscoloris could be recognised by its very dark, long wings and shiny orange-yellow mesosoma. Froggatt (1892) described the galls of P. acaciaediscoloris as formed at the inception of a leaf bud or new shoot. Sometimes, they would be little more than swellings at the base of the shoot. More often, they would be oval with three irregular horns formed from aborted leaf buds. Sometimes, P. acaciaediscoloris galls would be the target of inquilines of their own that caused the gall to degrade to a shapeless mass.

REFERENCES

Bouček, Z. 1988. Australian Chalcidoidea (Hymenoptera): A biosystematic revision of genera of fourteen families, with a reclassification of species. CAB International.

Froggatt, W. W. 1892. Notes on Australian Cynipidae, with descriptions of several new species. Proceedings of the Linnean Society of New South Wales, second series 7: 152–156.

The Chromeurytominae: Australo-Asian Mystery Wasps

One of the most diverse groups of micro-wasps is the Chalcidoidea, a bewildering array of intricate little jewels. A number of chalcidoid taxa have been extensively studied due to their roles as parasitoids of insect pests, but there are also many groups of chalcidoids that remain little known. One such group is the Chromeurytominae.

Male Chromeurytoma sp., copyright John Heraty.

The Chromeurytominae are a small group of chalcidoids primarily known from Australia, where they are represented by two genera, fourteen species of Chromeurytoma and the monotypic Asaphoideus niger (Bouček 1988). A single species, Pitayana coccorum, has also been described from Bangladesh (Bouček & Bhuiya 1990). Characteristic features include a relatively large subrectangular pronotum (the first segment of the thorax) and an antenna with six segments between the pedicel and the clava (the club). They are more or less shiny, often with a blue or green metallic gloss, and the gaster is fairly robust and does not collapse in preserved specimens. The affinities of the Chromeurytominae have been rather uncertain and the subfamily was only established by Bouček in 1988. Chromeurytoma itself was originally described in the family Eurytomidae, with which it shares the large pronotum. Other features suggest a relationship with the family Torymidae, such as an occipital carina (a ridge around the back of the head) and prominent cerci. Currently the Chromeurytominae are treated as part of the family Pteromalidae, which is not really saying too much. As our understanding of chalcidoid phylogeny has improved in recent years, it has largely confirmed what many workers had long suspected: that once you account for the other families, the Pteromalidae is pretty much just what's left over. Nevertheless, the broad-scale analysis of chalcidoids by Heraty et al. (2013) places the Chromeurytominae within a cluster of 'pteromalid' subfamilies, closer to the type subfamily Pteromalinae than to either the Eurytomidae or Torymidae.

The chromeurytomines are a bit of a mixed bag in terms of host species, but there is the common thread that their hosts are immobile or semi-sedentary plant-feeding insects. Pitayana coccorum attacks mealybugs and other soft scales, with multiple larvae potentially developing on a single host. Asaphoideus niger attacks the citrus leaf-miner Phyllocnistis citrella. The Chromeurytoma species are associated with galls on trees such as Eucalyptus; presumably they are parasites of the insects forming the galls.

REFERENCES

Bouček, Z. 1988. Australasian Chalcidoidea (Hymenoptera): A biosystematic revision of genera of fourteen families, with a reclassification of species. CAB International: Wallingford (UK).

Bouček, Z., & B. A. Bhuiya. 1990. A new genus and species of Pteromalidae (Hym.) attacking mealybugs and soft scales (Hom., Coccoidea) on guava in Bangladesh. Entomologist's Monthly Magazine 126: 231–235.

Heraty, J. M., R. A. Burks, A. Cruaud, G. A. P. Gibson, J. Liljeblad, J. Munro, J.-Y. Rasplus, G. Delvare, P. Janšta, A. Gumovsky, J. Huber, J. B. Woolley, L. Krogmann, S. Heydon, A. Polaszek, S. Schmidt, D. C. Darling, M. W. Gates, J. Mottern, E. Murray, A. D. Molin, S. Triapitsyn, H. Baur, J. D. Pinto, S. van Noort, J. George & M. Yoder. 2013. A phylogenetic analysis of the megadiverse Chalcidoidea (Hymenoptera). Cladistics 29: 466–542.

Sympiesis

Female Sympiesis, copyright Lyle J. Buss.

We often imagine that parasites select their hosts largely on the basis of type: one parasite prefers caterpillars, for instance, while another prefers flies. However, sometimes what is important is not so much what type of host a parasite attacks, as where they find it. The wasp in the photo above represents Sympiesis, a sizeable genus (the Universal Chalcidoidea Database lists over 130 species) of microscopic parasitoid wasps found worldwide. The majority of Sympiesis larvae attack the larvae of Lepidoptera, but others feed on the larvae of Diptera. A few have been recorded as hyperparasitoids, attacking the larvae of other parasitic wasps. The main thing that all hosts of Sympiesis have in common, though, is that they are all found in secluded, vegetative habitats: either mining in leaves, or in retreats formed by rolling or tying leaves (sometimes boring in stems). Depending on species, Sympiesis larvae may be either ectoparasites or endoparasites: those species feeding on leaf-rolling hosts tend to be ectoparasites, while those targeting leaf-miners are endoparasites (Miller 1970).

Sympiesis is a genus of the chalcid family Eulophidae. Eulophids used to be the subject of some disagreement between myself and a colleague of mine about their ease of recognition. Eulophids are a diverse group in appearance, coming in a bewildering array of shapes and colours. However, I have always maintained that they are nevertheless readily recognisable. Whatever their appearance, eulophids seem to always a distinctive stamp of 'eulophid-ness'. They tend to be slender, relatively soft-bodied wasps, often with a flat top to the gaster. Most identification guides will tell you to look out for their four-segmented tarsi (as opposed to the five-segmented tarsi of most other chalcid wasps); eulophid tarsi are rendered even more recognisable by the point that, though they have less segments than the tarsi of other chalcids, they are not any shorter so the individual tarsal segments are all relatively long. The features distinguishing Sympiesis from other eulophid genera are, of course, finer and require fairly close examination: notably, they have relatively few dorsal setae (only four on the scutellum) (Bouček 1988). As far as I know, they are mostly metallic green in coloration.

Male Sympiesis, showing branched antennae, from here.

As with many eulophids, males of Sympiesis usually differ from females in having long branches on the antennae. However, the first species of the genus to be described, the European Sympiesis sericeicornis, is distinctive in that these antennal branches are much reduced so that the males' antennae look little different from the females' (if you look very closely, they still have just a bit of a finger on each of the middle antennal segments). This led historically to a fair bit of confusion in the recognition of Sympiesis, with many species originally being placed in segregate genera (often with tongue-twistery compound names such as Asympiesiella or Sympiesonecremnus; thank you again, Alexandre Girault). Even now, the status of Sympiesis with regard to some related smaller genera could do with further investigation; we may yet see it grow again.

REFERENCES

Bouček, Z. 1988. Australasian Chalcidoidea (Hymenoptera): A biosystematic revision of genera of fourteen families, with a reclassification of species. CAB International: Wallingford (UK).

Miller, C. D. 1970. The Nearctic species of Pnigalio and Sympiesis (Hymenoptera: Eulophidae). Memoirs of the Entomological Society of Canada 102 (Suppl. S68): 5–121.

A New Short-horned Elasmus

Female Elasmus curticornis Gunawardene & Taylor 2012, newly out today!

I can now officially claim not to be a one-trick pony: my first non-harvestman academic paper has just been published. The paper, "New records of Elasmus (Hymenoptera, Eulophidae) species from Barrow Island, Western Australia", written with my co-worker Nihara Gunawardene, is freely available from the Journal of Hymenoptera Research.

Elasmus is a particularly attractive genus of chalcid micro-wasps that can be immediately distinguished from most other chalcids by their massively enlarged hind coxae, which are shaped like discs, and their long wedge-shaped wings. The Elasmus species of Australia were reviewed by Riek (1967), but most of them were known only from a small number of localities, mostly on the eastern side of the continent. In the course of going through material collected on Barrow Island, Nihara and I identified several species of Elasmus that had not been previously recorded from north-west Australia, and our new paper is mostly a record of those range extensions. Also, as most of the species had never actually been illustrated before, we provided extensive colour specimen photographs.

Elasmus ero emma, from Gunawardene & Taylor (2012).

Among the specimens, though, were a couple that we couldn't quite match up with any of the species in Riek's paper. They jumped between a few different identifications, but none of them really worked. So we had to broaden our comparisons: a bit of a daunting prospect, may I note, because Elasmus has over 200 species worldwide and I wasn't really keen on the idea of checking every single one of them to see whether they were the species we had on hand. As it turned out, I needn't have worried: the unusually short antennae of this species eliminated all but a few options. And after striking out those options as well, we prepared a description of a new species: Elasmus curticornis Gunawardene & Taylor 2012.

The species name means 'short-horned', in reference to the short antennae, and also in reference to one of the other similar species, E. brevicornis, which has an extensive distribution in Eurasia. I did spend a few days pondering whether our specimens might be slightly unusual examples of E. brevicornis: the most obvious difference between the two is that E. curticornis has a much more extensive area of orange on the gaster than has E. brevicornis. Eventually, we decided to go with declaring a new species, and at least none of the reviewers shot us down. I'm still keeping an eye out for more specimens to test our identification, but it doesn't seem to be a very abundant species so far.

Elasmus auratiscutellum, photographed by yours truly.

REFERENCES

Riek, E. F. 1967. Australian Hymenoptera Chalcidoidea family Eulophidae, subfamily Elasminae. Australian Journal of Zoology 15: 145–199.

Brachymeria perflavipes and Beyond

Male of a Brachymeria species, possibly B. hammari, photographed by John Hallmén.

For the subject of today's post, the wheel-spin brought up the parasitic wasp Brachymeria perflavipes. This is a species of the family Chalcididae described by our old friend Alexander Arsene Girault in 1913 from a female specimen collected in the Dandenong Mountains in Victoria, Australia. He described it under the name of Tumidicoxa flavipes; when that genus was later synonymised by Girault in 1915, the species name was replaced by perflavipes as there was already a Brachymeria flavipes. Girault's description of this species was, thankfully, more detailed than some of his later productions, though the species was not illustrated (not an uncommon state of affairs at the time) and, to the best of my knowledge, never has been. Girault (1915) later also recorded a male from Brisbane in Queensland, collected from a lepidopteran host. And that, except for an extra host record by Bouček (1988), is pretty much the sum total of our knowledge of this species.

Brachymeria calliphorae, a parasitoid of blow fly larvae, photographed by C. Bento.

Brachymeria is a large genus of Chalcididae, with over a hundred species worldwide. The Chalcididae are, in my opinion, one of the more instantly recognisable families of Chalcidoidea, with hind femora distinctly larger than those on other legs and with a ventral row of teeth. These large hind femora enable the wasp to jump, and I can confirm from personal observation that they are adept pingers. Most Chalcididae are black with coarse punctation over most of the body, though an interesting find on one of our recent field trips was a bright electric blue chalcidid. However, despite their recognisability, a recent major molecular phylogenetic analysis of Chalcidoidea failed to recover the Chalcididae as monophyletic. Brachymeria is distinguished from other chalcidid genera by features such as a short petiole, short postmarginal vein (but even shorter stigmal vein), distinct malar suture, and hind tibia with a distinct apical spine (Bouček 1988). Attempts have been made to subdivide Brachymeria into subgenera but, as we saw with Ormyrus, proposed subgeneric divisions have been based on species found within a restricted biogeographic area (e.g. Burks 1960; Bouček 1988) and a broad worldwide survey of Brachymeria still hasn't been conducted.

Brachymeria ovata on pupa of crow butterfly Euploea core, from Brisbane Insects.

Most Brachymeria species (including B. perflavipes) are parasitoids of lepidopteran pupae, but some attack dipteran larvae or pupae (more on that in a moment). Though more than one egg may be laid within a single pupa, Dowden (1935) found that in the case of B. intermedia, a parasitoid of the gypsy moth Lymantria dispar, only one wasp would develop to maturity in a single host. Indications are that the wasp larvae themselves kill any would-be competitors within the host: the mandibles of the larva are strongly sclerotised even in the first instar, and Dowden found that dead larvae dissected out from a host invariably bore signs of injury from the mandibles. Larvae dissected out from a host and placed together would quickly attack each other.

Diagram of distinguishing characters of Brachymeria intermedia vs B. compsilurae, from Dowden (1935).

Brachymeria species also include a good example of the potential importance of reliable taxonomic identifications. The aforementioned B. intermedia has been the subject of deliberate introductions outside its native range in attempts to control its host, the gypsy moth (or, more accurately, one of its wide range of hosts). Another Brachymeria species, B. compsilurae, closely resembles B. intermedia and also appears to lay its eggs in gypsy moths. However, the target of B. compsilurae is not the gypsy moth itself, but the larvae of tachinid flies that are parasitising the moth and are themselves used in its control. Programs to introduce B. intermedia to New England had to be careful to eliminate B. compsilurae (Burks 1960), whose release would have the opposite effect of that intended.

REFERENCES

Bouček, Z. 1988. Australasian Chalcidoidea (Hymenoptera): A biosystematic revision of genera of fourteen families, with a reclassification of species. CAB International: Wallingford (UK).

Burks, B. D. 1960. A Revision of the genus Brachymeria Westwood in America north of Mexico (Hymenoptera: Chalcididae). Transactions of the American Entomological Society 86 (3): 225-273.

Dowden, P. B. 1935. Brachymeria intermedia (Nees), a primary parasite, and B. compsilurae (Cwfd.), a secondary parasite, of the gypsy moth. Journal of Agricultural Research 50 (6): 495-523.

Girault, A. A. 1913. On several new genera and species of Australian Hymenoptera Chalcidoidea. Canadian Entomologist 45: 101-106.

Girault, A. A. 1915. Australian Hymenoptera Chalcidoidea—XIV. The family Chalcididae with descriptions of new genera and species. Memoirs of the Queensland Museum 4: 314-365.

Munro, J. B., J. M. Heraty, R. A. Burks, D. Hawks, J. Mottern, A. Cruaud, J.-Y. Rasplus & P. Jansta. 2011. A molecular phylogeny of the Chalcidoidea (Hymenoptera). PLoS One 6 (11): e27023. doi:10.1371/journal.pone.0027023.

Ormyrids: Attacking the Gall

Female of Ormyrus nitidulus, photographed by Penny Metal.

Everyone knows about God's supposed inordinate fondness for beetles, but it is my opinion that the true poster children for insect diversity should be the wasps. Wasps, admittedly, do not have as many described species as beetles (there are some who suspect that the actual number of species of wasp may eventually be higher, but that remains in the realm of the hypothetical). However, many species of beetle are very difficult to distinguish except by skilled specialists, being otherwise small, brown, and conservative. Wasps, on the other hand, come in a kaleidoscopic array of colours and shapes, such that even a novice may look at an array of wasps (see the top of this post, for instance) and be immediately struck by the disparity.

An unnamed species of Ormyrus, photographed by Simon van Noort.

The Chalcidoidea, commonly referred to as chalcids, are one of the largest subgroups of wasps, a clade of mostly small (often minute), mostly parasitoid wasps (some have larvae that feed on plants). Members of the Ormyridae, one of the commonly recognised families of chalcids, are generally about two to three millimetres long. Ormyrids are distinguished from other chalcids by their robust body form, with a strongly sclerotised gaster* (ormyrids and perilampids tend to look like steroid-abusing pteromalids). The segments of the gaster are usually ornamented by rows of coarse foveae (pits) that give it a distinctive rough appearance, though in some species these foveae are less obvious or are replaced by longitudinal ribs (Bouček 1988). Ormyrids are often recorded in association with plant galls, but are not gall-formers themselves: rather, they are parasites of the insect larvae that formed the galls (usually flies or other wasps). Some ormyrids are associated with figs and parasites of fig wasps.

*Wasp researchers generally refer to the sections of the body behind the head by terms such as 'mesosoma' and 'gaster' (or metasoma), rather than 'thorax' and 'abdomen'. This is because the section of the body that is the first segment of the abdomen in other insects has become the last segment of the mesosoma in Hymenoptera.

A female of Ormyrus on a knopper gall (a type of gall that develops when a developing acorn of the pedunculate oak Quercus robur is parasitised by the cynipid wasp Andricus quercuscalicis), photographed by Tristram Brelstaff.

There are about 125 known species of ormyrid (making this a quite small family by chalcid standards) according to the Universal Chalcidoidea Database (an absolutely wonderful resource). However, there isn't yet a really good classification system within the family. Ormyrids vary to a fair degree, particularly in the form of the antennae or the ornamentation of the gaster, but most authors have placed almost all species within the single genus Ormyrus. Attempts to subdivide this diverse group (for instance, that of Doğanlar, 1991, who recognised four genera of ormyrids with three subgenera within Cyrtosoma) have suffered from not considering the full range of ormyrid diversity. Some of the Australian forms referred to by Bouček (1988), for instance, may not be placeable in Doğanlar's system. Until an appropriately large-scale review is conducted, most authors will probably continue to recognise an all-purpose Ormyrus.

REFERENCES

Bouček, Z. 1988. Australasian Chalcidoidea (Hymenoptera): A biosystematic revision of genera of fourteen families, with a reclassification of species. CAB International: Wallingford (UK).

Doğanlar, M. 1991. Systematic positions of some taxa in Ormyridae and descriptions of a new species of Ormyrus from Turkey and a new genus in the family (Hymenoptera, Chalcidoidea). Türkiye Entomoloji Dergisi 15 (1): 1-13.