A re-examination of host relations in the Aphelinidae (Hymenoptera: Ghalcidoidea)

Host relations among the Aphelinidae (Hymenoptera: Chalcidoidea) are highly intriguing from an evolutionary view. Females are usually primary endoparasitoids of whiteflies or scale insects, whereas the development of the male is different and has been used for classification. In heteronomous hyperparasitoids (adelphoparasitoids or autoparasitoids) the male develops as a hyperparasitoid of conspecific females or other endoparasitoid species. We review the consequences of this mode of development which is unique because decisions concerning host selection are inextricably linked with progeny sex ratio. Autoparasitoid field sex ratios can fluctuate dramatically concurrent with changes in the relative availability of male and female hosts. A recent adaptive explanation for these sex ratios involves understanding the reproductive constraints acting on heteronomous parasitoids. Host relations in these parasitoids can show a degree of plasticity. We argue that in many instances autoparasitism may be facultative in nature and should not be used for classification. Heterotrophic parasitism, wherein the male develops as a primary parasitoid of lepidopterous eggs, has been poorly understood in the past due to uncertainties in reports of the biology and taxonomy of heterotrophic parasitoids. The host relations of this group are clarified.     

High nymphal host density and mortality negatively impact parasitoid complex during an insect herbivore outbreak

Insect herbivore outbreaks frequently occur and this may be due to factors that restrict top-down control by parasitoids, for example, host-parasitoid asynchrony, hyperparasitization, resource limitation and climate. Few studies have examined hostparasitoid density relationships during an in sect herbivore outbreak in a n atural ecosystem with diverse parasitoids. We studied parasitization patterns of Cardiaspina psyllids during an outbreak in a Eucalyptus woodland. First, we established the trophic roles of the parasitoids through a species-specific multiplex PCR approach on mummies from which parasitoids emerged. Then, we assessed host-parasitoid density relationships across three spatial scales (leaf, tree and site) over one yeas We detected four endoparasitoid species of the family Encyrtidae (Hymenoptera);two primary parasitoid and one heteronomous hyperparasitoid Psyllaephagus species (the latter with female development as a primary parasitoid and male development as a hyperparasitoid), and the hyperparasitoid Coccidoctonuspsyllae. Parasitoid development was host-synchronized, although synchrony between sites appeared constrained during winter (due to temperature differences). Parasitization was predominantly driven by one primary parasitoid species and was mostly inversely host-density dependent across the spatial scales. Hyperparasitization by C. psyllae was psyllid-density dependent at the site scale, however, this only impacted the rarer primary parasitoid. High larval parasitoid mortality due to density-dependent nymphal psyllid mortality (a consequence of resource limitation) compounded by a summer heat wave was incorporated in the assessment and resulted in density independence of host-parasitoid relationships. As such, high larval parasitoid mortality during insect herbivore outbreaks may contribute to the absence of host density-dependent parasitization during outbreak events.     

Endoparasitism in cereal aphids: molecular analysis of a whole parasitoid community

Insect parasitoids play a major role in terrestrial food webs as they are highly diverse, exploit a wide range of niches and are capable of affecting host population dynamics. Formidable difficulties are encountered when attempting to quantify host–parasitoid and parasitoid–parasitoid trophic links in diverse parasitoid communities. Here we present a DNA-based approach to effectively track trophic interactions within an aphid–parasitoid food web, targeting, for the first time, the whole community of parasitoids and hyperparasitods associated with a single host. Using highly specific and sensitive multiplex and singleplex polymerase chain reaction, endoparasitism in the grain aphid Sitobion avenae (F) by 11 parasitoid species was quantified. Out of 1061 aphids collected during 12 weeks in a wheat field, 18.9% were found to be parasitized. Parasitoids responded to the supply of aphids, with the proportion of aphids parasitized increasing monotonically with date, until the aphid population crashed. In addition to eight species of primary parasitoids, DNA from two hyperparasitoid species was detected within 4.1% of the screened aphids, with significant hyperparasitoid pressure on some parasitoid species. In 68.2% of the hyperparasitized aphids, identification of the primary parasitoid host was also possible, allowing us to track species-specific parasitoid-hyperparasitoid links. Nine combinations of primary parasitoids within a single host were found, but only 1.6% of all screened aphids were multiparasitized. The potential of this approach to parasitoid food web research is discussed.     

Classical biological control of ash whitefly: factors contributing to its success in California

The ash whitefly, Siphoninus phillyreae, invaded California in 1988 rapidly spreading throughout the state and infesting several species of ornamental trees and shrubs. Released Encarsia inaron rapidly established populations and spread throughout areas occupied by ash whitefly. We examined whether dispersal and overwintering ability could play a role in the extraordinary success of this parasitoid and we measured the impact of released parasitoids using a new method at a single location in northern California. The dispersal ability of E. inaron was examined by releasing two hundred and fifty adults 25 July 1991 into a single tree in a 1 ha pomegranate orchard near Brentwood, California. Based on yellow sticky card traps, the adult population spread at least 45 m from the release tree within 9 weeks of release. Over the same period of time the parasitoid population increased 64 fold. Impact was determined by measuring the number of ash whitefly adults produced in the absence and presence of E. inaron over 12 months. The production of whitefly adults, measured by the number of pupae entering the adult population, was reduced to very low numbers one year following the establishment of the parasitoid. Production of E. inaron adults increased rapidly, then decreased following the drop in production of ash whitefly adults. Duff of flowering pear trees was collected from under sample trees in mid winter to determine whether ash whitefly could survive on fallen leaves. Over fifty percent of adults emerged from whitefly pupae on leaves within 12 days of collection.     

Parasitoid-mediated transmission of an iridescent virus

We examined the interaction between an invertebrate iridescent virus (IIV) isolated from Spodoptera frugiperda (J.E. Smith) and the solitary ichneumonid endoparasitoid Eiphosoma vitticolle Cresson. In choice tests, parasitoids examined and stung significantly more virus infected than healthy larvae, apparently due to a lack of defense reaction in virus infected hosts. Parasitoid-mediated virus transmission was observed in 100% of the female parasitoids that stung a virus infected host in the laboratory. Each female parasitoid transmitted the virus to an average (+/-SE) of 3.7+/-0.3 larvae immediately after stinging an infected larva. Caged field experiments supported this result; virus transmission to healthy larvae only occurred in cages containing infected hosts (as inoculum) and parasitoids (as vectors). The virus was highly detrimental to parasitoid development because of premature host death and lethal infection of the developing endoparasitoid. Female parasitoids that emerged from virus infected hosts did not transmit the virus to healthy hosts. We suggest that the polyphagous habits of many noctuid parasitoids combined with the catholic host range of most IIVs may represent a mechanism for the transmission of IIVs between different host species in the field.     

Can hyperparasitoids cause large‐scale outbreaks of insect herbivores?

Synchronous population fluctuations occur in many species and have large economic impacts, but remain poorly understood. Dispersal, climate and natural enemies have been hypothesized to cause synchronous population fluctuations across large areas. For example, insect herbivores cause extensive forest defoliation and have many natural enemies, such as parasitoids, that may cause landscape‐scale changes in density. Between outbreaks, parasitoid‐caused mortality of hosts/herbivores is high, but it drops substantially during outbreak episodes. Because of their essential role in regulating herbivore populations, we need to include parasitoids in spatial modelling approaches to more effectively manage insect defoliation. However, classic host‐parasitoid population models predict parasitoid density, and parasitoid density is difficult to relate to host‐level observations of parasitoid‐caused mortality. We constructed a novel model to study how parasitoids affect insect outbreaks at the landscape scale. The model represents metacommunity dynamics, in which herbivore regulation, colonisation and extinction are driven by interactions with the forest, primary parasitoids and hyperparasitoids. The model suggests that parasitoid spatial dynamics can produce landscape‐scale outbreaks. Our results propose the testable prediction that hyperparasitoid prevalence should increase just before the onset of an outbreak because of hyperparasitoid overexploitation. If verified empirically, hyperparasitoid distribution could provide a biotic indicator that an outbreak will occur.     

The ectoparasitization of Coleophora alticolella (Lepidoptera) in relation to its aititudinal distribution

Abstract. 1. The parasitization of the larvae of Coleophora alticolella . feeding on Juncus squarrosus , was investigated at a series of altitudes from 15 to 520m above sea-level in northern England during 1977 and 1978.
2. Six species of primary parasitoid and one hyperparasitoid were reared from this host. Five of the primary parasitoids were ectophagous; only two specimens of the endoparasitoid, Gonotypus melanostoma , were reared.
3. All of the parasitoid species were recorded at 15 m but fewer at sites of higher altitude. Only one species, Scambus brevicomis , was recorded above 305 m, and none above 395 m. The hyperparasitoid, Tetrastichus endemus, was present only at 15 m.
4. Percentage parasitization was highest at 15 m; it was reduced from 51% to only 2% between 215 and 305 m in 1978. There was an increase in host density over this altitudinal range.
5. Three species, Scambus brevicomis. Elachertus olivaceus and Euderus viridis , accounted for most of the parasitization, but their relative proportions vaned at different altitudes.
6. The sex-ratios of the parasitoids reared from Coleophora alticolella ranged from 3.2% females for Scambus brevicomis , which is considered to also use larger hosts, to 99.4% females for Elachertus olivaceus , which develops by thelytokous parthenogenesis.
7. Euderus viridis and Scambus brevicomis started to attack the Coleophora alticolella larvae at a later date at 245 m than at 15 m, but attack by Elachertus olivacats was not delayed at the higher site.     

Sequential rapid adaptation of indigenous parasitoid wasps to the invasive butterfly Pieris brassicae

The introduction of a new species can change the characteristics of other species within a community. These changes may affect discontiguous trophic levels via adjacent trophic levels. The invasion of an exotic host species may provide the opportunity to observe the dynamics of changing interspecific interactions among parasitoids belonging to different trophic levels. The exotic large white butterfly Pieris brassicae invaded Hokkaido Island, Japan, and quickly spread throughout the island. Prior to the invasion, the small white butterfly P. rapae was the host of the primary parasitoid Cotesia glomerata, on which both the larval hyperparasitoid Baryscapus galactopus and the pupal hyperparasitoid Trichomalopsis apanteroctena depended. At the time of the invasion, C. glomerata generally laid eggs exclusively in P. rapae. During the five years following the invasion, however, the clutch size of C. glomerata in P. rapae gradually decreased, whereas the clutch size in P. brassicae increased. The field results corresponded well with laboratory experiments showing an increase in the rate of parasitism in P. brassicae. The host expansion of C. glomerata provided the two hyperparasitoids with an opportunity to choose between alternative hosts, that is, C. glomerata within P. brassicae and C. glomerata within P. rapae. Indeed, the pupal hyperparasitoid T. apanteroctena shifted its preference gradually to C. glomerata in P. brassicae, whereas the larval hyperparasitoid B. galactopus maintained a preference for C. glomerata in P. rapae. These changes in host preference may result from differential suitability of the two host types. The larval hyperparasitoid preferred C. glomerata within P. rapae to C. glomerata within P. brassicae, presumably because P. brassicae larvae attacked aggressively, thereby hindering the parasitization, whereas the pupal hyperparasitoid could take advantage of the competition-free resource by shifting its host preference. Consequently, the invasion of P. brassicae has changed the host use of the primary parasitoid C. glomerata and the pupal hyperparasitoid T. apanteroctena within a very short time.     

Impact of males on the progeny sex ratio of Campoletis chlorideae (Hym., Ichneumonidae), a parasitoid of Helicoverpa armigera (Hübner) (Lep., Noctuidae)

Abstract: The endoparasitoid Campoletis chlorideae and its host Helicoverpa armigera were reared in the laboratory to determine the progeny sex ratio of C. chlorideae in the presence and absence of male parasitoids. It was observed that the presence of male parasitoids affects the progeny sex ratio by decreasing the number of females in the offspring population, i.e. a significant increase in the proportion of males was observed in united sexes. Therefore, while constructing the life table of the parasitoid, the effect of presence of male parasitoids, should be considered together with other factors, which decreases the efficiency of the parasitoid.     

Description of a new Dendrocerus species (Hymenoptera: Megaspilidae) hyperparasitic on Stomaphis aphids and additional notes on their primary parasitoid, Protaphidius nawaii (Braconidae), and another hyperparasitoid, Euneura stomaphidis (Pteromalidae)

A food chain through three trophic levels on certain trees, Stomaphis aphids – a primary parasitoid, Protaphidius nawaii – a hyperparasitoid, Euneura stomaphidis , has been known in Japan. In the course of my survey on the seasonal prevalence of three Stomaphis species and their parasitoids from 1981–2007 in Kyoto, Japan, I found another hyperparasitoid, which I describe here as a new species, Dendrocerus stomaphis. From fragmentary observations, I inferred the life cycles of P. nawaii, E. stomaphidis and D. stomaphis . These three species have different strategies to escape ant aggression: P. nawaii by specific morphological and behavioral adaptation, E. stomaphidis by general anti-predator behavior like quick movement and D. stomaphis by a unique univoltine life cycle with the active phase from late autumn to early winter when the ants are less active.     

Rapid on-site identification of the biocontrol agent of the Asian chestnut gall wasp

In classical biocontrol programmes, a rapid and correct identification of the introduced antagonist is a key issue during both the release and establishment monitoring phases. It is often difficult to distinguish morphologically cryptic species or immature stages. An accurate diagnosis can now be provided by molecular diagnostic methods. Among the conventional and real-time PCR-based methods, loop-mediated isothermal amplification (LAMP) is a particularly suitable technique as it allows a rapid amplification of target DNA directly in the field. During the programme implemented in Italy against the Asian chestnut gall wasp (ACGW) Dryocosmus kuriphilus, we developed a real-time LAMP assay, combined with a simple DNA extraction method, for rapid in-field identification of larvae, pupae, and adults of the biocontrol agent, the parasitoid Torymus sinensis. Validation of the assay comprised adults as well as preimaginal stages of parasitoids obtained from ACGW galls collected from different localities. Results confirmed the effectiveness of the LAMP assay to rapidly and specifically identify the target parasitoid in the field. This assay will be a valuable tool for quick on-site checking of the parasitism rate.     

Effects of floral resources on the efficacy of a primary parasitoid and a facultative hyperparasitoid

Resources added to agroecosystems to enhance biological control are potentially available to multiple members of the resident insect community—not only the biological control agents for which the resources are intended. Many studies have examined the effects of sugar feeding on the efficacy of biological control agents. However, such information is lacking for other, interacting species such as facultative hyperparasitoids, which may contribute to pest suppression but can also interfere with introduced biological control agents. Under greenhouse conditions, we tested the direct effects of sugar and nectar provisioning on the longevity, host‐killing impact and offspring production of two pupal parasitoids associated with leek moth, Acrolepiopsis assectella: the introduced biological control agent, Diadromus pulchellus, and the native facultative hyperparasitoid, Conura albifrons. Adding sucrose, buckwheat or a combination of buckwheat and common vetch to a sugar‐deprived system (potted leek plants in cages) increased parasitoid longevity and resulted in higher leek moth parasitism and mortality compared to water or common vetch treatments. However, the two parasitoid species exhibited a distinct temporal response to the treatments, likely influenced by differences in their life histories. This study provides insight into how integrating conservation biological control techniques could affect the success of a classical biological control programme.     

Interspecific competition between Diadegma semiclausum Hellen and Diadegma mollipla (Holmgren), parasitoids of the diamondback moth, Plutella xylostella (L), feeding on a new host plant

Interspecific competition between an introduced parasitoid species aimed at controlling a herbivorous pest species and a native parasitoid parasitising the same host may influence the success of classical biological control programmes. In Kenya, interspecific competition between an introduced and a local parasitoid on two diamondback moth populations (DBM, Plutella xylostella) was investigated on two different host plants. We tested simultaneous and delayed competition of the local parasitoid Diadegma mollipla Holmgren and its exotic congenus D. semiclausum Hellen on a newly aquired DBM host plant (snowpea) in the laboratory. Under simultaneous competition, D. mollipla produced more progeny than D. semiclausum on snowpea. A head start of D. Mollipla, of four and eight hours before its congenus was introduced, resulted in a similar number of progeny of both species. In delayed competition (time intervals of 24 h, 48 h and 72 h), progeny production was similar for both parasitoids when the time interval was 24 h, irrespective of which species parasitized first. More progeny was produced by the species which attacked first, when the time interval was greater than 24 h, although it was only significant at 72 h. Competitive abilites of both parasitoids on the new host plant differed largely between laboratory and semi-field conditions. The influence of two host plants (snowpea and cabbage) on competition was studied in the greenhouse with different host and parasitoid densities. Parasitism levels of D. semiclausum were significantly higher than those of D. mollipla, regardless of host plant, host and parasitoid densities, but progeny production of D. mollipla on snowpea was still slightly higher than on cabbage. As compared to the confinement of parasitoids and larvae to small containers, D. mollipla parasitized very few larvae in the cages. Competitive ability of the two parasitoid species tested was influenced both by the density of the searching females and by parameters related to either the host plant and/or the herbivorous hosts.     

Native primary parasitoids and hyperparasitoids attacking an invasive aphid Uroleucon nigrotuberculatum in Japan

Since the invasion of Uroleucon nigrotuberculatum from North America we searched for parasitoids of this aphid on Solidago altissima in Japan to determine what species of native parasitoids attack the newly invasive aphid. We found three primary parasitoid species: Ephedrus plagiator and Praon yomenae (Braconidae, Aphidiinae) and Aphelinus albipodus (Aphelinidae). We also found eight hyperparasitoid species: Syrphophagus sp. (Encyrtidae), Dendrocerus carpenteri (Megaspilidae), Asaphes suspensus (Pteromalidae) and Pachyneuron aphidis (Pteromalidae) through both E. plagiator and A. albipodus; Phaenoglyphis villosa (Figitidae, Charipinae), Aprostocetus sp. (Eulophidae, Tetrastichinae) and D. laticeps through E. plagiator, and Alloxysta sp. nr brevis (Figitidae, Charipinae) through A. albipodus. Uroleucon nigrotuberculatum is usually attacked by rather polyphagous primary parasitoids, E. plagiator and A. albipodus, in Japan, where an oligophagous parasitoid specialized to allied aphid species is probably absent. The hyperparasitoid community of U. nigrotuberculatum is common to those of the aphids occurring in open field‐type habitats in Japan.     

Parasitoid assemblages of soybean defoliator Lepidoptera in north-western Buenos Aires province, Argentina

1 The objective of this study was to examine the structure of parasitoid assemblages attacking soybean defoliator Lepidoptera in north-western Buenos Aires province, Argentina. 2 Through larvae sampling from 1992 to 1996, and laboratory data plus the known biology of the species recorded during the study, parasitoid species composition, richness, number of guilds per host species, and levels of parasitism, were determined. 3 Total species richness was 23, all species were primary endoparasitoids, and categorized as koinobionts. They belonged to Hymenoptera (11 species) and Diptera (12 species). Rachiplusia nu (Noctuidae) hosted the highest number of parasitoid species. 4 Four parasitoid guilds were recorded for R. nu (egg–prepupal endoparasitoid, early larval endoparasitoid, late larval endoparasitoid, and larval–pupal endoparasitoid); two for Spilosoma virginica (Arctiidae) (late larval endoparasitoid and larval–pupal endoparasitoid) and Colias lesbia (Pieridae) (early larval endoparasitoid and larval–pupal endoparasitoid); and one for Anticarsia gemmatalis (Noctuidae) and Loxostege biffidalis (Pyralidae) (early larval endoparasitoid). 5 Only four out of seven potential parasitoid guilds defined for Lepidoptera were recorded in a soybean agroecosystem located in north-western Buenos Aires province, suggesting that potential host niches were not totally utilized. 6 Soybean lepidopteran defoliators supported a mean species richness of 4.8 parasitoids, a result that is similar to that reported for exophytic hosts in the Neotropics. 7 Parasitoid assemblages from each host differed in species composition, richness, number of guilds, and levels of parasitism.     

Autoparasitism, interference, and parasitoid-pest population dynamics

Autoparasitoids ("heteronomous hyperparasitoids") are parasitoids that lay female eggs on homopteran hosts and male eggs on juvenile parasitoids of either the same species or another species. Males develop as hyperparasitoids and eventually kill the juvenile parasitoid. We present a series of stage-structured models that investigate the effects of autoparasitism on population dynamics. Autoparasitism causes density-dependent mortality on juvenile parasitoids and therefore has a stabilizing effect. This also leads to an increase in host population abundance. In most cases an autoparasitoid leads to higher host equilibrium densities than a comparable primary parasitoid (except when the primary parasitoid is arrhenotokous (sexual) and the autoparasitoid has a low preference for attacking parasitized hosts or can attack the parasitized host for only a small portion of its development). When male autoparasitoids are followed explicitly in the models, mate limitation reduces the stabilizing effect of autoparasitism and leads to a further increase in host abundance. Coexistence of an autoparasitoid with a nonprimary parasitoid or second autoparasitoid is possible when the level of conspecific autoparasitism is greater than the level of heterospecific autoparasitism. When an autoparasitoid coexists with a primary parasitoid, the resulting host density is always greater than that with only the primary parasitoid. Therefore, autoparasitoids have the potential to disrupt control achieved by primary parasitoids. When two autoparasitoids coexist, the resulting host density is always lower than that attained by either autoparasitoid alone. The effects of autoparasitism are compared with those of other forms of interference competition.     

Aphid parasitoids detect that an alien plant was present nearby during their development

Abstract Glasshouse and laboratory experiments were conducted with Aphidius rhopalosiphi parasitizing Metopolophium dirhodum to test whether the presence in the same environment of another plant species (tomato or marigold) induced aerial allelopathy in wheat plants.
Emerging parasitoids were offered the choice of two odours of wheat plants in an olfactometer – wheat grown alone and wheat grown in the presence of the second plant species.
Female parasitoids responded more strongly than males, but both sexes preferred the odour of wheat grown in the environment in which they had developed (i.e. with or without the second plant species present).
Moving the tomato plants from the cages where they had been placed with wheat to the cages that had previously had wheat alone just before the aphid mummy formed showed that the odour environment the emerging parasitoid responded to was that of the mummy rather than that of the developing parasitoid larva. From this it was concluded that the parasitoids in the olfactometer were responding not only to wheat volatiles, but also to tomato volatiles absorbed on the surface of the mummy and encountered by the emerging adult parasitoid.
By excising the parasitoid pupa from the mummy formed on one wheat cultivar, and offering the emerging parasitoids the surface of aphid mummies formed on a different cultivar, it was confirmed that olfactometer responses represented a learning of the chemistry of the surface of the mummy encountered at or after emergence.     

Effects of dietary nicotine on the development of an insect herbivore, its parasitoid and secondary hyperparasitoid over four trophic levels

Abstract.  1. Allelochemicals in herbivore diet are known to affect the development of higher trophic levels, such as parasitoids and predators.
2. This study examines how differing levels of nicotine affects the development of a herbivore, its parasitoid and secondary hyperparasitoid over four trophic levels. Separate cohorts of the herbivore, Manduca sexta , were fed on artificial diets containing 0.0, 0.1, and 0.5% wet weights of nicotine. Some of the larvae in each cohort were separately parasitised in the first (L1) and third (L3) instars by the gregarious endoparasitoid, Cotesia congregata . Newly emerged parasitoid cocoons were, in turn, parasitised by the hyperparasitoid, Lysibia nana .
3. Pupal mass in M . sexta was negatively correlated with nicotine concentrations in the artificial diet, although larval development time was unaffected.
4. Hyperparasitoid survival was highest when there were low levels of nicotine in the diet of M . sexta . Cocoon mass in C . congregata and adult mass in L . nana were mostly affected by nicotine levels in host diet when L1 M. sexta larvae were parasitised. The effects were slightly stronger on L . nana than on C . congregata , indicating the presence of both qualitative and quantitative effects of nicotine concentration on both species.
5. The results suggest that allelochemicals in herbivore diet can have both direct and indirect effects on the performance of higher trophic levels. However, in multitrophic interactions these effects can vary with the stage of the herbivore attacked by the primary parasitoid, as well as with the strategy employed by the herbivore to deal with plant phytotoxins.     

Host selection by the heteronomous hyperparasitoid Encarsia pergandiella: multiple-choice tests using Bemisia argentifolii as primary host

The ovipositional patterns of the heteronomous hyperparasitoid Encarsia pergandiella Howard (Hymenoptera: Aphelinidae) in the presence of its primary host Bemisia argentifolii Bellows & Perring (Hemiptera: Aleyrodidae), and in the presence or absence of conspecific and heterospecific secondary hosts (Encarsia formosa Gahan andEretmocerus mundus Mercet; Hymenoptera: Aphelinidae) were examined to assess host species preferences. Host preferences by heteronomous hyperparasitoids may affect the relative abundance of co-occurring parasitoid species and may influence host population suppression by the parasitoid community. Four combinations of hosts were tested: (1) B. argentifolii, E. mundus, and E. formosa, (2) B. argentifolii, E. formosa, and E. pergandiella, (3) B. argentifolii, E. mundus, and E. pergandiella, and, (4) B. argentifolii, E. mundus, E. formosa, and E. pergandiella. Arrays of hosts (24) were constructed in Petri dishes using leaf disks, each bearing one host. Thirty arrays of each host combination were exposed to single females for 6 h. All hosts were dissected to determine number of eggs per host. Encarsia pergandiella parasitized E. formosa hosts as frequently as E. mundus hosts. However, E. pergandiella parasitized either of these heterospecific hosts more frequently than conspecific hosts in treatments including two secondary host species. When a third parasitoid species was included in host arrays, E. pergandiella parasitized conspecific hosts as frequently as heterospecific hosts. Developmental stage of the hosts did not significantly influence host species selection by E. pergandiella. Our results indicate that host selection and oviposition by heteronomous hyperparasitoids like E. pergandiella, vary with the composition of hosts available for parasitization, and suggest a preference for heterospecific over conspecific secondary hosts.     

Flexible larval development and the timing of destructive feeding by a solitary endoparasitoid: an optimal foraging problem in evolutionary perspective

1. In studying the evolution of life-history strategies in parasitoids, considerable attention has been paid to the relationship between host quality and parasitoid fitness. Various workers have reported that host quality influences parasitoid size, development time, and survival. Because body size is frequently correlated with fecundity, longevity, and host-finding ability in parasitoids, this parameter is usually considered to be the main target of selection. 2. In koinobiont parasitoids that consume the entire host before pupation, adult parasitoid size and development time are often strongly correlated with host size at the time when it is developmentally arrested through destructive feeding by the parasitoid larva. 3. Here, a mathematical model is proposed to describe the larval feeding behaviour of the solitary koinobiont endoparasitoid Venturia canescens in four larval stadia of its host Plodia interpunctella. In particular, the model describes how adult size, represented by an exponential growth rate, and development time are traded off when the parasitoid develops in nutritionally suboptimal second stadium hosts. 4. Using a graphical model, the different conditions faced by V. canescens during development in various host species of greatly differing mass are illustrated. 5. It is argued that the relative importance of size and development time on parasitoid fitness is determined by ecological and biological characteristics of both host and parasitoid, and it is suggested that there may be correlations between life-history traits and host-utilisation strategies among koinobionts.