Evolution of life-history timing in a leafmining moth: Phenotypic selection in patches with manipulated development time

Phyllonorycter mespilella (Hübner) is a leafmining moth with two stages of larval development: the initial sap-feeding (SF) stage followed by the tissue-feeding (TF) stage. Phenotypic selection by parasitoids on the duration of the SF stage (SF duration) was measured in artificial patches of larvae placed in the field during the diapausing generation. Pretreatment of larvae with different photoperiods allowed creation of patches that varied in the time-course of appearance of TF larvae. The shorter the photoperiod pretreatment, the sooner TF larvae tended to appear. Some patches were left exposed and others were caged to exclude parasitoids. Positive directional selection on SF duration was detected in exposed patches, and no selection was detected in caged patches. Directional selection in exposed patches was caused by both parasitoid oviposition and other unidentified sources of mortality. The other sources of mortality may have included host feeding by parasitoids on TF larvae. A larger proportion of parasitoid eggs were oviposited on TF larvae in patches where TF larvae appeared the earliest, but this variation in parasitoid oviposition did not result in significant differences in directional selection intensity among patches with early, intermediate and late appearance of TF larvae. Although the general form of the fitness function was very similar when compared among patch types, no significant directional selection could be detected in patches where TF larvae appeared late, and the causes of directional selection appeared to vary between patches where TF larvae appeared at early and intermediate dates.     

Size-selective oviposition by parasitoids and the evolution of life-history timing in hosts: fixed preferences vs frequency-dependent host selection

The influence of size-selective oviposition behaviour by parasitoids on the evolution of life-history timing in their hosts was examined using an optimization model of a two-stage life history similar to a genetic algorithm. Host populations with varying durations of early-larval development were subjected to selection in scenarios where parasitoids had fixed preferences for oviposition on late-stage larvae, or those where parasitoid attack was dependent on the relative frequencies of the two life stages present in the population. Fixed preference for oviposition on late-stage larvae caused positive directional selection on the duration of early-larval development. Surviving individuals remained for as long as possible in the first stage of development in order to avoid parasitoid attack. Frequency-dependent parasitoid attack, in contrast, caused maintenance of variation in the duration of early-larval development. The influence of the fitness payoffs of different life stages on the plasticity of size-selective oviposition behaviour is discussed, as are possible implications of the model results for parasitoid-host population dynamics.     

VARIABLE SELECTION ON EUROSTA'S GALL SIZE,I: THE EXTENT AND NATURE OF VARIATION IN PHENOTYPIC SELECTION

Natural fluctuations in environmental conditions are likely to induce variation in the intensity or direction of natural selection. A long-term study of the insect, Eurosta solidaginins Fitch (Diptera; Tephritidae), which induces stem galls on the perennial herb Solidago altissima (Asteraceae) was performed to explore the patterns of variation in phenotypic selection. The intensity of selection imposed by parasitoids and predators on gallmaking larvae, for gall size, was measured across 16 populations over the course of 4 generations, for a total of 64 population-generations. Directional selection was quantified by i, the selection intensity, and variance selection by j‘, a measure of the intensity of selection on phenotypic variance. Size-dependent attack by parasitoids caused upward directional selection (mean i_p = 0.42; SE = 0.023), while size-dependent bird attack favored larvae that induced smaller galls (mean i_b = -0.07; SE = 0.013. The mean net directional selection intensity was 0.35 (SE = 0.030), which indicates that insects inducing larger galls are generally favored by selection. The opposing patterns of size-dependent attack resulted in stabilizing selection in half the population generations, with an overall average. j‘ of -0.11 (SE = 0.078). The magnitude of directional selection was strongly influenced by the population mean gall size and weakly by the optimal gall size. The intensity of variance selection was strongly influenced by the shape of the fitness function, with sigmoidal and Gaussian-like shapes causing greater depletion of phenotypic variance.     

VARIABLE SELECTION ON EUROSTA'S GALL SIZE. II. A PATH ANALYSIS OF THE ECOLOGICAL FACTORS BEHIND SELECTION

We examined phenotypic selection exerted by natural enemies on the gall-making fly Eurosta solidaginis in an extensive field study of 16 populations, spanning four generations. Gall-makers that induce small galls are vulnerable to the attack of Eurytoma gigantea. This imposes upward directional selection on gall size. Insectivorous birds, predominantly the downy woodpecker, are more likely to attack larvae that induce large galls than small ones, and this imposes downward directional selection. We used path analysis to explore the relative contributions of these natural enemies to the net directional selection on gall size. The path models further examined several ecological factors that influence selection intensity through their effects on parasitoid and bird attack rates. Net directional selection varied more strongly with E. gigantea attack than bird attack. Competitive interactions among birds and the three parasitoid species, including E. gigantea, were evidenced by low winter bird attack rates in fields where a high proportion of galls contained the overwintering parasitoids. Eurytoma gigantea attack was heavier in fields where mean gall size was small and bird attack heavier in fields where mean gall size was large. Neither birds nor E. gigantea showed simple density-dependent attack. Data suggested a form of frequency-dependent attack by birds but not by E. gigantea.     

Interaction of a solitary larval endoparasitoid, Microplitis mediator, with its host, Mamestra brassicae: host acceptance and host suitability

Abstract:  Microplitis mediator (Haliday) (Hym., Braconidae) is an important parasitoid of early instar larvae of the European cabbage moth, Mamestra brassicae L. (Lep., Noctuidae). In the laboratory, we examined attack responses of female M. mediator to the first three larval instars of M. brassicae . Females were presented with M. brassicae larvae either one individual at a time in a no-choice experiment, or three individuals, one from each instar, simultaneously in a choice experiment. Whether or not there was choice, naïve female parasitoids attacked a high proportion of larvae and did not discriminate among instars. In the no-choice experiment, attacked larvae were reared, and parasitoid cocoons were produced from about 76% of larvae attacked as first and second instars, but from only 19% of larvae attacked as third instars. Dissections of attacked larvae from the choice experiment showed that about 79% of attacks on first and second instars resulted in oviposition compared with only 49% for third instars. When given choice, frequency and number of attacks on first instar larvae increased with increasing parasitoid experience. Our results suggest that first and second instar larvae of M. brassicae are suitable hosts for M. mediator , but that third instar larvae are suboptimal both because oviposition attempts were frequently unsuccessful and because immature parasitoids failed to complete development. Nevertheless, naïve attacking parasitoids exhibited minimal discrimination among instars, although experienced parasitoids most frequently attacked first instar larvae. The host selection behaviour of M. mediator is discussed in the context of optimal foraging theory and implications for biological control.     

Variation in selection pressures on the goldenrod gall fly and the competitive interactions of its natural enemies

Summary Larvae of the tephritid fly Eurosta solidaginis induce ball-shaped galls on the stem of tall goldenrod, Solidago altissima. Survival probability depends on gall size; in small galls the larva is vulnerable to parasitoid oviposition, whereas larvae in large galls are more frequently eaten by avian predators. Fly populations from 20 natural old fields in central Pennsylvania were monitored in 1983 and 1984 to examine the distribution of the selection intensity imposed by natural enemies, the parasitoids Eurytoma gigantea and E. obtusiventris, the inquiline Mordellistena unicolor, and the predatory birds Dendrocopus pubescens and Parus atricapillus. Mordellistena and E. obtusiventris are able to attack galls of all diameters while E. gigantea and the predatory birds preferentially assaulted small and large diameter galls, respectively. Eurosta in intermediate sized galls had the highest survivorship, hence selection had a stabilizing component. However, parasitoid attack was more frequent than bird attack, and the two did not exactly balance, thus there was also a directional component. The mean directional selection intensity on gall size was 0.21 standard deviations of the mean, indicating that larger gall size was favored. Interactions among the insect members of the Eurosta natural enemy guild are complex and frequent.     

Reproductive biology and small-scale rearing of cherry bark tortrix and its candidate biological control agent

Abstract:  The larval parasitoid, Campoplex dubitator (Hym.,: Ichneumonidae), is under consideration as an agent for classical biological control of cherry bark tortrix (CBT), Enarmonia formosana (Lep.,: Tortricidae), in North America. A comprehensive risk analysis of the candidate agent will require prosperous cultures of both the pest and its parasitoid. We present a rearing method for small-scale production of both species using a bean-based artificial diet, with additional information on the reproductive biology of C. dubitator . Based on estimated survivorship probabilities, a CBT egg had a 70% chance of developing completely to the adult stage under this system. The success of parasitism, however, was very dependent on the instar of the CBT host larva at the time of oviposition. All parasitised first instar larvae died shortly after the attack, rendering them unsuitable for oviposition, while 50% of parasitised second instar larvae died prematurely. In contrast, early mortality was 15–30% for larvae parasitised in the third to fifth instars. Regardless of the instar at oviposition, approximately 90% of the surviving hosts yielded parasitoids, showing a high acceptance by C. dubitator of second to fifth instars for oviposition.     

Weak parasitoid-mediated apparent competition between two Phyllonorycter (Lepidoptera: Gracillariidae) leaf miner species on a deciduous oak Quercus dentata

Parasitoid assemblages and the rates of parasitism on tissue-feeding larvae of two Phyllonorycter leaf miner species, P. persimilis and P. leucocorona , were studied from the autumn generation in 2002 to the summer generation in 2005 to understand whether parasitoids mediate interactions between the two leaf miner species. Fourteen species of parasitoids emerged from P. persimilis and 11 emerged from P. leucocorona . The parasitism rate was high: 24.1–92.6% in P. persimilis and 58.9–81.7% in P. leucocorona . Thus, parasitism was a major mortality factor in the present Phyllonorycter species. The parasitoid composition was distinctly different between the two host species, although most parasitoids were able to parasitize both leaf miner species. The analysis based on the quantitative parasitoid overlap revealed that the present parasitoids could mediate interactions between the present leaf miner species, but their effects would be weak. This is attributable to parasitoid's preferential uses of either of the leaf miners as a host.     

SELECTION ON HERBIVORE LIFE-HISTORY TRAITS BY THE FIRST AND THIRD TROPHIC LEVELS: THE DEVIL AND THE DEEP BLUE SEA REVISITED

Abstract The timing of life‐history events in insects can have important consequences for both survival and reproduction. For insect herbivores with complex life histories, selection is predicted to favor those combinations of traits that increase the size at metamorphosis while minimizing the risk of mortality from natural enemies. Studies quantifying selection on life‐history traits in natural insect herbivore populations, however, have been rare. The purpose of this study was to measure phenotypic selection imposed by elements of the first and third trophic levels on variation in two life‐history traits, the timing of egg hatch and pupal mass, in a population of oak‐feeding caterpillars, Psilocorsis quercicella (Lepidoptera: Oecophoridae). Larvae were collected from the field throughout each of two generations per year for three years and reared to determine the effects of the date of egg hatch on both the risk of attack from parasitoids and the pupal mass of the survivors. The direction and strength of phenotypic selection attributed to aspects of the first and third trophic levels, as well as their combined effects, on the date of egg hatch was measured for each of the six generations. Heritabilities of and genetic correlations between pupal mass and the date of adult emergence from diapause (the life‐history trait expected to have the largest influence on the timing of egg hatch, and thus larval development) were estimated from laboratory matings. In four of the six generations examined, significant directional selection attributed to the first trophic level was detected, always favoring early‐hatching cohorts predicted to experience higher leaf quality than late‐hatching cohorts. Directional phenotypic selection by the third trophic level was detected in only one of three years, and in that year the direction of selection was in opposite directions during the two successive generations. The combined effect of selection by both trophic levels indicated that the third trophic level acted to either reduce or enhance the more predictable pattern of selection attributed to the first trophic level. In addition, I found evidence of truncation selection acting to increase the mean and decrease the variance of pupal mass during the pupa‐adult transition in the laboratory. Pupal mass and diapause duration were found to vary significantly among full‐sibling families; upper bounds for heritability estimates were 0.57 and 0.30, respectively. Furthermore, these two traits were found to be positively genetically correlated (families with larger pupae had longer diapause durations), resulting in a fitness trade‐off, because larger pupae enjoy higher survival through metamorphosis and female fecundity but emerge later, when average leaf quality for offspring is generally poorer.     

Variations in parasitism in sympatric populations of three invasive leaf miners

Abstract:  Parasitism was investigated in sympatric populations of three invasive gracillariid leaf miners in Europe, Cameraria ohridella , Phyllonorycter robiniella and Phyllonorycter platani to test the hypotheses that C. ohridella is less heavily parasitized by native parasitoids and attacked by fewer species than the two other invasive species. In all regions investigated, C. ohridella showed a lower parasitism rate, and its parasitoid complex was poorer in species than those of either Phyllonorycter spp. Comparisons were made between sympatric populations of C. ohridella on its main host tree, Aesculus hippocastanum , and an occasional host, Acer pseudoplatanus . Parasitism rates were similarly low and composed of the same parasitoid species on both trees. In contrast, a sympatric population of Phyllonorycter geniculella , a native species mining A. pseudoplatanus , was heavily parasitized by a totally different parasitoid complex. These results suggest that the low parasitism in C. ohridella by native polyphagous leaf miner parasitoids is due neither to its host tree, nor to a problem of synchronization between the phenology of the pest and that of its parasitoids. Instead, it probably results from the inability of the native parasitoids to locate, attack, or develop on a new host that does not have any native congener in Europe.     

Scarlet gilia resistance to insect herbivory: the effects of early season browsing,plant apparency,and phytochemistry on patterns of seed fly attack

Pollen-limited plants are confronted with a difficult tradeoff because they must present showy floral displays to attract pollinators and yet must also minimize their apparency to herbivores. In these systems, traits that increase pollinator visitation may also increase herbivore oviposition and overall plant resistance may therefore be constrained to evolve largely as a correlated response to selection on plant apparency or vigor. We used a family-structured quantitative genetic experiment to evaluate the importance of ungulate browsing, flowering date and plant height (traits that are related to overall vigor), and variation in a putative phytochemical defense (cucurbitacin production) on patterns of seed fly attack in a scarlet gilia population. We found significant genetic variation in the amount of insect damage plants experience in the field, providing evidence that resistance may evolve. In addition, we found that browsing reduced seed fly attack and that oviposition is strongly related to plant size and flowering date; large, early flowering plants experience high attack. In addition, we found that high cucurbitacin production was correlated with low seed fly damage, although this effect was relatively weak.We found directional selection on final plant height and flowering date; tall, early flowering plants had the highest reproductive success. In addition, we found negative directional selection on cucurbitacin production, which may indicate a high cost of cucurbitacin or other functions of this phytochemical. Although seed fly herbivory arguably decreases plant fitness, we found an unexpected positive relationship between damage and fitness. A negative relationship between fitness and damage may be masked in this system through strong positive indirect correlations between patterns of damage and levels of pollinator visitation. Finally, we found significant genetic variation in flowering date, plant height, and cucurbitacin production. Resistance to seed flies may evolve in this population, but largely as a non-adaptive correlated response to selection on overall plant vigor. Phytochemicals may play a more important role in defense in years with high seed fly attack, or when pollen-limitation is less severe.Co-ordinating editor: J. Tuomi     

Does mother really know best? Oviposition preference reduces reproductive performance in the generalist parasitoid Aphidius ervi

The reproductive success of female parasitoids is dependent on their ability to accurately assess the suitability of a host for larval development. For generalist parasitoids, which utilize a broad range of species and instars as hosts, a set of assessment criteria determines whether a host is accepted or rejected. The suitability of a host, however, can only be imperfectly assessed by the female parasitoid, which can result in the selection of lesser quality hosts for oviposition. In this study we explored the disparity between host quality and host preference using the generalist koinobiotic parasitoid Aphidius ervi Haliday (Hymenoptera: Aphidiidae) and the host Aulacorthum solani (Harris) (Homoptera: Aphididae), the foxglove aphid. The second instar hosts produced the highest level of reproductive success, while third and fourth instars resulted in a substantially reduced reproductive performance. When given a choice of host instars, parasitoids preferred the older hosts for oviposition disregarding their reduced suitability for larval development. Results are discussed in context of mechanisms involved in A. ervi host selection and biases in the criteria used to assess hosts that may arise when parasitoids transfer host species between generations.     

Biosteres longicaudatus: Developmental dependence on host (Anastrepha suspensa) physiology

Larvae of Anastrepha suspensa that were in the first day of the third instar were parasitized by females of the solitary endoparasitoid, Biosteres longicaudatus. At the end of the 6-hr oviposition period, larvae were ligated posterior to the ring gland so that some larvae had parasitoids anterior to the ligature while in others, the parasitoids were in the abdomen, posterior to the ligature. Ninety-two percent of the parasitoids anterior to the ligature hatched to the first through third instars. Parasitoids posterior to the ligature had a 75% egg hatch to the first instar only. No larval molts to the second or subsequent instars occurred in these parasitoids. Upon parabiosis to 3-day-old, unparasitized host pupae, the ligated larvae pupated and 97% of the first-instar parasitoids in these parabiosed larval abdomens molted to the second instar. Newly laid parasitoid eggs transplanted to 3-day-old pupal hosts had less than one-third of the egg hatch of those transplanted to first-day third-instar hosts. The data implicate the physiological state of the host (vis-a-vis pupation and associated events) as being an important factor in the development of the endoparasitoid.     

Factors affecting host selection in an insect host–parasitoid interaction

1. Many parasitoids can develop successfully in different stages of the same host but the costs of parasitism may vary between the stages. The stage of host attacked has generally been determined when there is no choice, giving a misleading impression of host selection or preference.
2. The rate of parasitism by a solitary endoparasitoid, Venturia canescens , of each larval stage of the Indian meal moth, Plodia interpunctella , was examined with and without a host refuge from parasitism. In addition, when given a choice of host stages, with and without a refuge, the influence of parasitoid age on host selection was examined.
3. Wasps were able to parasitize all except the first instar, but second instars experienced significantly reduced parasitism, in both refuge treatments, compared with third to fifth instars. Whilst parasitoid emergence was always significantly less when all host stages had a refuge, the reduction was only marginally significant when second instars were attacked.
4. When given a choice of second- and fifth-instar larvae, wasps consistently parasitized more fifth instars, both with and without a refuge. Moreover, significantly fewer second-instar larvae were parasitized in the presence of fifth instars than when presented alone to the wasps. This pattern of parasitism was unaffected by the increasing age of the parasitoids.
5. Host selection by V. canescens is discussed in terms of host–parasitoid population dynamics and structure.     

Seasonal phenology and stage-specific parasitism of the apple ermine moth, Yponomeuta malinellus Zeller, in Korea

The apple ermine moth, Yponomeuta malinellus Zeller (Lepidoptera: Yponomeutidae), is a tent caterpillar that feeds on Malus spp. in Korea. Populations of the moth in native areas appeared to be regulated by the assemblage of parasitoids. Phenological associations between host stages and parasitoids, susceptible stage(s) of the host for each parasitoid, and stage‐specific parasitism were studied. The egg larval parasitoid Ageniaspis fuscicollis (Dalman) had highest parasitism of first instar larvae (24%), with 14% parasitism of other larval stages. Dolichogenidea delecta (Haliday) was recovered from all larval instars with the highest parasitism rate of second instar larvae (20.1%), followed by 19.9% parasitism of mid‐larval hosts. Herpestomus brunicornis Gravenhorst was reared from second instar larvae through to pupal collection, and had the highest parasitism rate (29.9%) at the pupal stage. The larval pupal parasitoid Zenillia dolosa (Meigen) was recovered from mid‐larval to pupal stages with the highest parasitism rate (5.5%) occurring in third to fourth instar larvae. The host stages for developing A. fuscicollis completely overlap with those of D. delecta, and with those of H. brunicornis to some degree. A statistically significant negative correlation exists between A. fuscicollis and these dominant parasitoids, indicating competitive interaction within the host.     

Effects of Bacillus thuringiensis on Adults of Cotesia plutellae (Hymenoptera: Braconidae), a Parasitoid of the Diamondback Moth, Plutella xylostella (Lepidoptera: Plutellidae)

To complement existing information on the mortality of larvae of the wasp Cotesia plutellae attacking moth caterpillars infected with Bacillus thuringiensis (Bt) we tested the direct and indirect effects of the bacterium on adult wasp longevity and oviposition behaviour. In one experiment with female parasitoids, mean longevity (SEM) was not significantly different between females exposed to Bt (1.98 +/- 0.08 days) and those not exposed (2.18 +/- 0.13 days). In a second experiment with both males and females, Bt treatment did not significantly effect either male or female parasitoids exposed to Bt. To observe the possible effects of Bt on oviposition behaviour of C. plutellae each of 10 females were given five larvae that had been treated with Bt and five untreated larvae at the same time. All parasitoids were observed to make oviposition attempts in both untreated and treated larvae. Upon dissection of the host larvae, one or more C. plutellae eggs were found in each of the larvae in which a parasitoid attempted oviposition. There was no effect of Bt treatment on parasitoid oviposition. The mean number of ovipositions in treated larvae (4.3 +/- 0.3) was not significantly different from untreated larvae (4.7 +/- 0.2).     

Choosy egg parasitoids: Specificity of oviposition-induced pine volatiles exploited by an egg parasitoid of pine sawflies

Generalist parasitoids are well‐known to be able to cope with the high genotypic and phenotypic plasticity of plant volatiles by learning odours during their host encounters. In contrast, specialised parasitoids often respond innately to host‐specific cues. Previous studies have shown that females of the specialised egg parasitoid Chrysonotomyia ruforum Krausse (Hymenoptera: Eulophidae) are attracted to volatiles from Pinus sylvestris L. induced by the egg deposition of its host Diprion pini L. (Hymenoptera: Diprionidae), when they have previously experienced pine twigs with host eggs. In this study we investigated by olfactometer bioassays how specifically C. ruforum responded to oviposition‐induced plant volatiles. Furthermore, we studied whether parasitoids show an innate response to oviposition‐induced pine volatiles. Naïve parasitoids were not attracted to oviposition‐induced pine volatiles. The attractiveness of volatiles from pines carrying eggs was shown to be specific for the pine and herbivore species, respectively (species specificity). We also tested whether not only oviposition, but also larval feeding, induces attractive volatiles (developmental stage specificity). The feeding of D. pini larvae did not induce the emission of P. sylvestris volatiles attractive to the egg parasitoid. Our results show that a specialist egg parasitoid does not innately show a positive response to oviposition‐induced plant volatiles, but needs to learn them. Furthermore, the results show that C. ruforum as a specialist does not learn a wide range of volatiles as some generalists do, but instead learns only a very specific oviposition‐induced plant volatile pattern, i.e., a pattern induced by the most preferred host species laying eggs on the most preferred food plant.     

Seasonal population changes of five parasitoids attacking the scale insect Nipponaclerda biwakoensis on the common reed, with special reference to predation by wintering birds

Seasonal changes in the abundance of five species of hymenopterous parasitoids (four species of Encyrtidae and one species of Eulophidae) attacking the scale insect Nipponaclerda biwakoensis on the common reed were investigated for 2 years in Lake Biwa, with special reference to predation by the reed bunting, Emberiza schoeniclus, during winter. The scales settled on reed shoot stems under sheath leaves, passing through three discrete generations per year. The abundance of adult female scales increased exponentially from July (first generation) to December (third generation). Adult female scales of the third generation overwintered on reed shoots. During winter, female scale abundance dramatically declined, whereas the number of predation marks made by reed buntings using their bills on reed sheath leaves increased. The generations of all five parasitoids were synchronized with the host scale generations, and the five parasitoids overwintered as larvae inside the scale bodies. The abundance of parasitized scales and parasitoid adults emerging from the scales also increased from July to December, but greatly decreased during winter. The overall parasitism rate of the female scales remained at relatively low levels (less than 40%) throughout the year, including before and after winter. A bird exclusion experiment revealed that the dramatic winter decrease of the abundance of the scale and its five parasitoids was due to intensive and non‐selective predation by the buntings on unparasitized and parasitized scales. Additionally, the proportion of immature parasitoids removed by birds varied between the five parasitoid species. Thus, seasonal population changes of the five scale parasitoids are considerably affected by bird predation on overwintering immature parasitoids.     

Regulation of host diapause by an insect parasitoid

Abstract. 1. The interaction between larval development and parasitism by the braconid wasp Cotesia koebelei (Riley), was investigated in a population of the butterfly Euphydryas editha (Boisduval) (Nymphalidae). In this population, the butterfly host has an obligatory overwintering larval diapause.
2. It was found that E. editha larvae harbouring parasitoids were more likely to pass through an extra feeding instar before entering diapause than were non-parasitized conspecifics.
3. In addition, some individuals that were experimentally exposed to multiple parasitoid attacks bypassed diapause completely; these larvae passed through five or six feeding instars, reaching sizes typical of final instar post-diapause larvae.
4. The observed effect of superparasitism occurred regardless of whether the host larvae subsequently produced mature parasitoids, suggesting that parasitoid attack is sufficient to invoke the response.
5. It is proposed that the parasitoid C.koebelei regulates the number of pre-diapause feeding instars of its insect host E. editha, and that some component of the female venom, injected at oviposition, is responsible for this regulation.     

Host selection and survival of the parasitoidEncarsia formosa on greenhouse whitefly,Trialeurodes vaporariorum, in the presence of hosts infected with the fungusAschersonia aleyrodis

Successful control of greenhouse whitefly may be achieved by complementary activity of the parasitoidEncarsia formosa and the fungusAschersonia aleyrodis. One way to obtain an additive mortality effect of both entomopathogen and parasitoid would be achieved by the selection of healthy hosts by the parasitoid and rejection of fungus-infected hosts. Third and fourth instar larvae ofTrialeurodes vaporariorum which had been treated with a spore suspension ofA. aleyrodis 0, 4, 7, 10 or 14 days beforehand, were presented to female parasitoids. The parasitoids adopted the oviposition posture on untreated hosts as well as on treated hosts, irrespective of the different stages of infection in the hosts. However, significantly more hosts were parasitized byE. formosa in the control treatment than in the fungal treatment. The parasitoids offered treated hosts, showed rejection behaviour after probing on hosts showing detectable signs of infection (containing hyphal bodies or mycelium in the haemolymph). For instance, when hosts were offered seven days after spore treatment, the parasitoids showed an oviposition posture on a total of 83 (95.4%) out of 87 infected larvae, but laid only 4 eggs (4.6%). In contrast, on 48 (94.1%) out of 51 noninfected (or showing no detectable signs of infection) hosts an oviposition posture was adopted and 40 eggs (78.4%) were found after dissection. When infected hosts were encountered the oviposition posture lasted less than 1′40″ while rejection of non-infected hosts occurred after more than 1′40″. Other experiments were carried out offering treated hosts for 24 h to the parasitoids. The hosts were dissected afterwards. Again, significantly more eggs were laid in the non-infected hosts. When hosts were parasitized shortly after fungal spore treatment they were colonized by the fungus and the parasitoids did not develop. Transmission of the entomopathogen after probing infected hosts was observed to a limited extent. In conclusion,A. aleyrodis andE. formosa can be used together in a glasshouse situation. The parasitoid will be most effective when introduced more than seven days after application ofA. aleyrodis, because from that time onwards it is able to detect and reject fungus-infected hosts.