The four major biological strategies of ichneumonoid parasitoids, koinobiont and idiobiont, ecto-and endoparasitism, are discussed and the evolutionary radiations of the two families Ichneumonidae and Braconidae compared in an attempt to relate differences in patterns of host utilization to differences in evolutionary history. The most primitive members of both families are idiobiont ectoparasitoids of hosts concealed in plant tissue. Idiobiont ectoparasitic braconids are all still primarily associated with such hosts, but idiobiont ectoparasitic ichneumonids have radiated to attack hosts in other situations, such as in aculeate nests or in cocoons. A shift in emphasis between the behavioural steps, host habitat location and host location, is envisaged as being important in such evolutionary change. Idiobiont endoparasitism is postulated as having arisen amongst ectoparasitoids attacking cocooned hosts, as an adaptation that allows them to exploit pupae and puparia in relatively exposed positions; it is a fairly common strategy in the Ichneumonidae, but virtually unknown in the Braconidae. Koinobiosis is perceived as having evolved in association with hosts which feed in a relatively weakly concealed position, but pupate in a more secluded and safe location. The strategy is advantageous as it allows a parasitoid to oviposit on an easily discoverable host, but to use the host's pupation concealment to complete its own development. The evolution of koinobiosis has allowed parasitoids to exploit hosts that feed in exposed positions, and to attack hosts at a younger and numerically more common stage in the host's life cycle. Koinobiont ectoparasitism is envisaged, in some braconid and ichneumonid groups, to occupy an evolutionary transitional position between idiobiosis and endoparasitic koinobiosis; only in the Ichneumonidae have large radiations of koinobiont ectoparasitoids occurred. Endoparasitic koinobiosis is hypothesized as having arisen in the Braconidae in association with lepidopterous/coleopterous hosts, whilst in the major lineage of endoparasitic koinobiont ichneumonids, this habit is hypothesized as having arisen in association with symphytan hosts. The great majority of braconids are koinobiont endoparasitoids, but only about 50% of the Ichneumonidae have this habit. Very few koinobiont braconids develop as endoparasitoids of hymenopterous hosts, although many endoparasitic ichneumonids attack Hymenoptera. However, lineages of the Braconidae have radiated to exploit adult insects and exopterygote nymphs; ichneumonids do not utilize such hosts.

Evolutionary patterns of host utilization by ichneumonoid parasitoids (Hymenoptera: Ichneumonidae and Braconidae)*

Many turf managers prefer to control foliage- and root-feeding pests with the same application, so-called multiple-targeting, using a single broad-spectrum insecticide or a premix product containing two or more active ingredients. We compared the impact of a neonicotinoid (clothianidin), a premix (clothianidin + bifenthrin), and an anthranilic diamide (chlorantraniliprole), the main insecticide classes used for multiple targeting, on four species of beneficial insects: Harpalus pennsylvanicus, an omnivorous ground beetle, Tiphia vernalis, an ectoparasitoid of scarab grubs, Copidosoma bakeri, a polyembryonic endoparasitoid of black cutworms, and Bombus impatiens, a native bumble bee. Ground beetles that ingested food treated with clothianidin or the premix suffered high mortality, as did C. bakeri wasps exposed to dry residues of those insecticides. Exposure to those insecticides on potted turf cores reduced parasitism by T. vernalis. Bumble bee colonies confined to forage on white clover (Trifolium repens L.) in weedy turf that had been treated with clothianidin or the premix had reduced numbers of workers, honey pots, and immature bees. Premix residues incapacitated H. pennsylvanicus and C. bakeri slightly faster than clothianidin alone, but otherwise we detected no synergistic or additive effects. Chlorantraniliprole had no apparent adverse effects on any of the beneficial species. Implications for controlling turf pests with least disruption of non-target invertebrates are discussed.

Impacts of a neonicotinoid, neonicotinoid-pyrethroid premix, and anthranilic diamide insecticide on four species of turf-inhabiting beneficial insects.

During an outbreak of army cutworm in southern Alberta in the spring of 1990, the overall incidence of parasitism by the polyembryonic parasitoid, Copidosoma bakeri (Howard), was 61% in samples from seven fields. The incidence of parasitism in samples of army cutworms collected on five dates from a single location, during the spring of 1991, increased from about 20% in the early samples to about 50% in the later samples. Cutworms parasitized by C. bakeri feed for a longer time than unparasitized ones; therefore estimates of the incidence of parasitism by C. bakeri, based on samples of late-instar cutworms, are misleadingly high. Parasitized cutworms also grow considerably larger than unparasitized ones and may have a supernumerary instar. Larger hosts support larger broods of C. bakeri and apparently a successful strategy of C. bakeri is to prolong host development so as to maximize an acquired resource. Because cutworms parasitized by C. bakeri feed more and longer than unparasitized cutworms, a high rate of parasitism can exacerbate crop damage and complicate control recommendations. The life cycles of army cutworm and C. bakeri are asynchronous and it is likely that high rates of parasitism are dependent on the presence of intermediary hosts.

Parasitism of the army cutworm, euxoa auxiliaris (grt.) (lepidoptera: noctuidae), by copidosoma bakeri (howard) (hymenoptera: encyrtidae) and effect on crop damage

Polyembryonic parasitoids producing single-sex broods of clonal offspring provide an unusually clear window into the classic tradeoff between the number and size of offspring. We conducted a laboratory study of the encyrtid parasitoid Copidosoma bakeri parasitizing the noctuid Agrotis ipsilon to examine the way that size and number of offspring tradeoff in broods of each sex and to determine how the fit between host and parasitoid brood is achieved. We found that brood mass (wasp body massbrood size) was proportional to host mass, independent of brood sex, indicating a tight fit between brood and host and ensuring a sizenumber tradeoff. By correcting brood size and body mass of each brood for host mass, we demonstrated the expected inverse relationship between wasp variables. We postulated that the wasp brood might achieve the fit to the host by (1) adjusting brood size based on information available early in host development before and during division of the embryo, (2) manipulating host size late in host development after completion of embryo division, or (3) simply adjusting individual wasp mass to fill the host. We evaluated host responses to parasitismand correlations between brood size and host growth early and late in developmentfor broods of each sex. The data are consistent with adjustment of brood size to the amount of host growth early in host development and with manipulation of host mass late in host development. The tight link between host mass and brood mass also suggests a final adjustment by parasitoid growth to achieve complete filling. Within the tight fit, female broods were smaller but contained larger individuals than male broods. The sex-specific balance point of the tradeoff and sex differences in balancing mechanisms and responses to host size suggest different selection pressures on each sex requiring future investigation.

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A sex‐specific size–number tradeoff in clonal broods

Larvae of bertha armyworm, Mamestra configurata Walk., collected in 1972–75 on rapeseed in the prairie provinces of Canada (predominantly in Manitoba), were killed by one virus, one fungus, one microsporidium, seven insect parasites, and by other factors. Mortality from disease was about 24% in 1972 when bertha armyworm was abundant, but much lower in succeeding years after the host’s population density declined. The most important hymenopterous parasite, Banchus fiavescens Cress., was relatively rare in 1972 but subsequently killed over 40% of the host larvae each year. A tachinid, Athrycia cinerea (Coq.), killed over 20% of the mature bertha armyworms in each generation. Each of the other five parasite species was associated infrequently with M . configurata . Details of the biology of each pathogen and parasite are presented. About 12% of the mature host larvae died because they failed to empty the gut, with subsequent penetration of the gut wall by bacteria, or because they did not form a morphologically-sound pupa. Pupae of M . configurata died in the soil following penetration of the gut wall by bacteria, as a result of soil cultivation, and from low temperatures in winter. No pathogens or parasites were recorded in field-collected pupae, though an ichneumonid, Ichneumon canadensis Cress., parasitized pupae in a field cage and may occasionally do so in natural conditions. No pathogens or parasites of bertha armyworm eggs were recorded. A preliminary survival model for bertha armyworm, based on estimates of mortality from various causes identified in this study, is presented. The possibility of preventing or reducing future oubreaks of M . configurata by colonizing additional parasite or pathogen species in the prairie provinces is discussed.

A. C. Schaaf

Papers

The parasitoid complex of Euxoa ochrogaster (Guenee) (Lepidoptera: Noctuidae)