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.

Wasps that don't Give to a Fig

The strategies employed by flowering plants to draw in their pollinators are many and varied. Some have entered into exclusive partnerships, contriving methods by which their rewards are shared with a single animal species and hence presumably increasing the likelihood of that species visiting them. Once remarkable example of such a partnership is found among the figs. To the casual observer, fig trees might appear to never produce flowers. However, immature figs are in fact closed inflorescences called syconia with the flowers produced on the inside of the fig, never exposed to the outside world. The only way for pollinators to reach the fig flowers is through a tiny hole or ostiole at the fig's apex. This ostiole is used by the fig's pollinators, tiny female wasps of the chalcidoid family Agaonidae, who enter the fig in search of places to lay their eggs. The wasp herself does not leave the fig again after laying but her eggs and larvae develop within galls inside the fig, feeding on the tissue of the fig itself. After developing into wingless males and winged females, the next generation of fig wasps mates within the syconium; pollination of the fig tree occurs through the young females leaving the fig to find their own laying places and carrying pollen as they do so.

Female Idarnes nr flavicollis, a typical late-laying sycophagine, copyright Sergio Jansen Gonzalez.

The fig benefits by having an exclusive pollinator, the fig wasp benefits by having a ready-made nursery for its offspring. However, all that tasty fig tissue is bound to prove attractive to others who would circumvent the standard contract. Another group of chalcidoid wasps, the subfamily Sycophaginae, includes prime examples of such freeloaders. Like the true pollinating fig wasps, these non-pollinating fig wasps develop in galls within fig syconia. However, instead of entering the fig through the ostiole, most sycophagines use their ovipositor to pierce the fig's outer skin and lay from outside. In some sycophagines, the ovipositor is relatively short and thick; these species lay their eggs when the fig is only just beginning to develop. In others, the ovipositor is longer and slender, longer in fact than the rest of the wasp, and oviposition happens later when the fig has grown to a larger size. In some of these later-arriving forms, the ovipositing female lays into a gall already induced by the fig's actual pollinator (how she finds it from outside the fig, I have no idea), and as well as feeding on the gall, her larva will eventually feed on the pollinator larva. There are also some sycophagines that enter the syconium through the ostiole and oviposit internally; I haven't been able to find whether these species may function as true pollinators.

Female Sycophaga ovipositing on Ficus sur, copyright JMK.

As noted above, pollinating agaonids exhibit strong sexual dimorphism with only the females having wings, and males never escaping the host syconium. The wingless males cannot be easily recognised as belonging to the same species as the females; indeed, if one does not already know what they are, they can barely even be recognised as wasps. In sycophagines, matters are a bit more complicated. In some species, males are wingless and highly modified as in pollinating fig wasps. In others, males are winged and similar in appearance to females. And in still others, wingless and winged males are both present within a single species. I don't know what determines whether a given larva of these species develops wings or not; both forms may develop within the same syconium. It has been suggested that the presence of the two forms is related to conflicting pressures of gene flow vs speed. Winged males that can look for mates outside the parent syconium have a better chance of finding mates outside the pool of their own siblings, thus avoiding the risk of inbreeding. However, wingless males can mate with females immediately after they emerge within the syconium (if not before, as I'll explain shortly), in which case the winged males may simply find themselves too late to the party. Certainly there is a correlation between winglessness and the size of broods. Early-ovipositing species, which tend to produce smaller broods because the younger host syconium offers less space for egg-laying, are more likely to have winged males whereas males of later-ovipositing species are more likely to be wingless (Cruaud et al. 2011).

Male Apocryptophagus, copyright Centre for Biodiversity Genomics.

Males of the genera Sycophaga and Apocryptophagus (which Cruaud et al., 2011, suggested should probably be synonymised) are invariably wingless and have elongate, flattened, extensible gasters. The terminal pair of spiracles on the abdomen have the surrounding peritremes (supporting rings) extended into a pair of long filaments. In the host figs of these genera, the interior of the syconium becomes filled with liquid after being pollinated by its associated agaonids; the liquid is resorbed when the pollinators emerge. Nevertheless, there are advantages for the sycophagines in emerging before the pollinators: not only could the interior of the syconium become rather crowded, males of some agaonid species have enlarged mandibles that they may use to dispatch any interlopers. So instead of waiting for the syconial fluid to drain away, the male Sycophaga cuts a small opening slit in his gall (too narrow for the surrounding fluid to seep in) through which he partially emerges into the central cavity. The peritremal filaments are used to anchor the end of his gaster within his original gall so that he can continue to breathe from the air-pocket inside it while he stretches out in search of another gall containing a female. When he finds one, he will cut into it in the same way that he cut out of his own, then release the end of his gaster from its anchor-point and quickly slip into the gall of his intended. After a brief mating, he can repeat the process, this time using the female's gall as his air-tank (Ramírez 1996–1997). The female presumably emerges once the syconial fluid is gone.

Male Apocryptophagus emerging from a gall containing a female, from Ramírez (1996–1997).

There have been various viewpoints about the relationships of Sycophaginae to other chalcidoids. Some authors have included almost all the fig-associated wasps in the Agaonidae, whether pollinators or not. Others have restricted the Agaonidae to the true pollinators and classified non-pollinating fig wasps such as the Sycophaginae with the poorly defined family Pteromalidae. An analysis of chalcidoid relationships by Heraty et al. (2013) identified Sycophaginae as a sister group to Agaonidae sensu stricto (while placing other groups of non-pollinating fig wasps elsewhere on the tree). This might lead one to consider the possibility that the gall-making habit seen in both Sycophaginae and pollinating Agaonidae pre-dates the evolution of the wasp-fig relationship as pollinators. Perhaps the evolution of the syconium allowed figs to convert gall-makers that had previously been parasites into partners.

REFERENCES

Cruaud, A., R. Jabbour-Zahab, G. Genson, F. Kjellberg, N. Kobmoo, S. van Noort, Yang D.-R., Peng Y.-Q., R. Ubaidillah, P. E. Hanson, O. Santos-Mattos, F. H. A. Farache, R. A. S. Pereira, C. Kerdelhué & J.-Y. Rasplus. 2011. Phylogeny and evolution of life-history strategies in the Sycophaginae non-pollinating fig wasps (Hymenoptera, Chalcidoidea). BMC Evolutionary Biology 11: 178.

Ramírez, W. 1996–1997. Breathing adaptations of males in fig gall flowers (Hymenoptera: Agaonidae). Revista de Biologia Tropical 44 (3)–45 (1): 277–282.