General Biology

Since Hymenoptera is a very large order, it is not surprising that a considerable number of biologies and life history strategies are exhibited by its various taxa. Broadly speaking, the basal lineages are phytophagous as larvae, feeding both ecto- and endophytically on a large range of herbs, shrubs, and trees; few tropical pergid sawflies, even feed on slime molds! The great majority of the remaining species are either parasitoids of other insects or predators of insects (e.g., the yellow-jackets or social wasps, which are members of the Vespidae) or spiders. However, among the higher taxa, there have also been several reversals to phytophagy, especially through the formation of galls on plants (cecidogenesis). The bees (Apidae) and one other, small tropical group, the Masarinae within the Vespidae, have evolved to make use of pollen and nectar as a larval food source.

Like other holometabolous groups, hymenopterans primitively pass through generally five instars, though the number of instars is typically smaller in endoparasitic taxa, and in one such genus there seems to be just a single instar. The final instars of hymenopterans are rather morphologically conservative, with most sawflies having rather caterpillar-like larvae with well-developed true legs and variously developed prolegs on several of the abdominal segments. Adoption of an endophytic way of life by cephoid sawflies and wood wasps was accompanied by a reduction in the prolegs and more generally by reduction of sensory structures. Final instar apocritan wasp larvae are all quite similar and are termed hymenopteriform. They are superficially rather maggotlike in that they lack legs and other processes and often have a rather reduced head. However, many endoparasitoids have highly bizarre, first instars characteristic of their particular families, and for which a variety of specific terms have been coined.

The pupal stage of hymenopterans is exarate; that is, the antennae, legs, and wings are free from the body (in contrast to the Lepidoptera, e.g., in which these components are fused with the body). The pupae tend to be rather delicate and are easily damaged. All sawflies and most members of the Ichneumonoidea + Aculeata clade produce a silken cocoon to protect the pupa. Most of the other parasitic taxa do not, however, probably because they pupate within the host remains or, if they pupate externally, do so in a location where the pupa is likely to be protected by the surroundings, such as within a leaf mine, gall, or wood boring.

Key Features in Hymenoptera Evolution

Given the huge size of the order, it is interesting to consider what features have enabled hymenopterans to be so successful in terms of both individuals and total number of species. Most attention has focused on a small number of features such as selection of oviposition site, modification of that site, the use of venoms, and the evolution of the thin wasp waist, all of which are discussed in this article. In addition, the unusual form of sex determination mechanism, haplodiploidy, may have been particularly important in the evolution of sociality. It is likely, however that few of these traits have operated in isolation, and it is the interactions of these and other factors that have been important. Thus, for example, evolution of sociality may have been facilitated by the sex determination mechanism but also requires the abilities to remember where the nest is, to recognize nestmates, and to be able to defend the nest.

Several studies have emphasized that the success of the Hymenoptera has probably been a consequence of the general tendency of these insects to provide their offspring with particularly nutritious food sources, and when necessary (and that has been often) to modify poorer foods to better ones. Although this may be most familiar in terms of the provisioning of larvae in the nests by the social wasps and bees, such

TABLE I Most Recent Classification of the Sawflies and Wood Wasps



Described extant species


Xyeloidea Pamphilioidea

Xyelidae Pamphiliidae


Tenthredinoidea Argidae
















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