Performance of moth larvae on birch in relation to altitude, climate, host quality and parasitoids

We studied topographical and year-to-year variation in the performance (pupal weights, survival) and larval parasitism of Epirrita autumnata larvae feeding on mountain birch in northernmost Finland in 1993–1996. We found differences in both food plant quality and parasitism between sites ranging from 80 m to 320 m above sea level. Variation in food plant quality had particularly marked effects on larval survival. The advanced phenology of the birches in relation to the start of the larval period reduced pupal weights. Parasitism rates were different between years and between sites. The clearest site differences were in the proportions of different parasitoid species: Eulophus larvarum was most abundant at the lowest-altitude sites, and Cotesia jucunda at the highest. Differences in the performance of E. autumnata were related to temperature conditions: at higher temperatures, survival and the egg production index were lower, and larval parasitism was higher than at lower temperatures. The higher parasitism at higher temperatures was probably due to greater parasitoid activity during warmer days. In the comparison of different sources of spatial and annual variation in the performance of E. autumnata, the most important factor appeared to be egg mortality related to minimum winter temperature, followed by parasitism and, finally, the variation in food plant quality. If, as predicted, the climate gradually warms up, the effects of warmer summers on the outbreaks of E. autumnata suggest a decrease in outbreak intensity. Received: 4 January 1999 / Accepted: 22 March 1999     

Host-feeding and oviposition strategies of parasitoids in relation to host stage

Until now, mathematical models of parasitoid-host interactions have not incorporated the tendency for destructively host-feeding parasitoids to partition their feeding and oviposition behaviour in relation to different host stages. A literature survey reveals a trend for female parasitoids to feed preferentially or exclusively on earlier host stages and to oviposit preferentially or exclusively in/or later ones. We explore the relative advantages to host-feeding parasitoids of a number of possible host stage selection strategies. We develop hypotheses, formalizing and testing them using modifications to our earlier simulation model of host-feeding strategies (Jervis and Kidd, 1986). We conclude from our modelling that the advantage to be gained from feeding on early host stages and ovipositing in late ones is likely to be associated with: 1) reduced handling times when feeding on early stage hosts; 2) reduced wastage of progeny from mortality factors other than host-feeding by the parent parasitoid, achieved by confining oviposition to late host stages; and 3) reduced probability of progeny mortality resulting from the parent's host-feeding activities.     

The influence of oviposition phenology on survival in host races of Eurosta solidaginis

Phenological differences between host plants can contribute to allochronic isolation of host races of phytophagous insects. Host races of the gallmaking fly, Eurosta solidaginis, that live on different species of goldenrod (Solidago altissima and S. gigantea) exhibit different emergence phenologies. These differences could result from adaptation to corresponding phenological differences between the hosts in periods of optimal suitability for gall formation and survival to adulthood. In order to test this, some flies of each host race were allowed to emerge naturally while the emergence times of others were manipulated to correspond to the emergence and oviposition periods of the other host race. Percent gall formation and survival to adulthood were examined for three oviposition periods: the peak time of emergence and oviposition of the earlier-emerging host race (that from S. gigantea), that of the later-emerging host race (that from S. altissima), and a week after the peak emergence of the host race from S. altissima. Flies of both host races were allowed to ovipuncture plants of the appropriate species during each of these periods. Plant relative growth rates were measured during each of these periods. The experiment was repeated twice over a two-year period. Relative growth rates of both host species were highest during the earliest oviposition period (the period during which the host race from S. gigantea normally emerges). Percent gall formation was significantly correlated with plant relative growth rate, but the coefficient of determination was low. In both years of the study, percent gall formation of both host races was highest during the earliest oviposition period (the period during which the host race from S. gigantea normally oviposits). Likewise, percent survival to adulthood in both host races was highest during the earliest oviposition period. There was no significant effect of oviposition period on the percent of larval death due to parasitism by Eurytoma gigantea or predation by Mordellistena unicolor. These results suggest that the host race from S. altissima does not emerge at the time that its host is optimally suited for gall formation or survival to adulthood. Therefore, differences in emergence phenologies do not appear to be due to corresponding phenological differences between the host species in suitability for gall formation or survival to adulthood.     

Temporal and spatial variation of larval parasitism in non-outbreaking populations of a folivorous moth

In order to assess the role of parasitoids in the regulation of non-outbreaking populations of Epirrita autumnata, a geometrid lepidopteran with outbreaking populations in northern Europe, we examined the temporal and spatial variation of larval parasitism in southwestern Finland during 6 successive years. The study was carried out on two spatial scales, among trees within sites of about 1 ha and among sites separated by distances of 2–10 km, using experimental and observational approaches respectively. The overall percent parasitism was independent of host density on both spatial scales, while temporally it fluctuated only little. Of the two main parasitoids, the commoner one, Protapanteles immunis, showed a variable response to host density on the larger spatial scale and negative density dependence on the smaller scale. Temporally, parasitism caused by this species was independent of host density. Another parasitoid, Phobocampe bicingulata, showed positive density dependence on the smaller spatial scale and had a variable response on the larger scale, but exhibited negative density dependence over time. The results of this study caution against drawing conclusions concerning population regulation on the grounds of spatial density dependence alone. Larval parasitoids apparently do not maintain low densities in the E. autumnata populations studied. However, they may suppress E. autumnata densities to a level low enough for density-dependent mortality factor(s) to become regulating. Among other mortality factors of E. autumnata, pupal predation has been found to be temporally positively density-dependent. Received: 19 October 1999 / Accepted: 10 January 2000     

Host-feeding and oviposition by parasitoids in relation to host stage: Consequences for parasitoid-host population dynamics

Among parasitoids which host-feed destructively, there is a tendency for females to partition their feeding and oviposition behaviour in relation to different host stages, feeding preferentially or exclusively on earlier host stages and ovipositing preferentially or exclusively in (or on) later ones. We explored the dynamic implications of this behaviour for parasitoid-host population dynamics, using modifications of the age-structured simulation models of Kidd and Jervis (1989, 1991). Using the new versions of the models, we compared the situation where parasitoids practice host stage discrimination with respect to feeding and oviposition, with the situation where they do not. Additionally, we examined the effects of host stage discrimination on populations by (a) having generations either discrete or overlapping, (b) varying initial age structure, (c) having varying degrees of density dependence acting on host adult mortality, and (d) varying parasitoid develoment times in relation to the length of host development. With either discrete or overlapping generations of the host population, a reduction in the parasitoid development time had a destabilizing influence on the parasitoid-host population interaction. With discrete generations stage discrimination had no effect on the risk of extinction, irrespective of either the degree of density dependence acting on the host population, or the initial age structure of the host population. When parasitoid search was uncoupled from the insect's adult energy requirements, the interaction was always unstable. With continuous generations, stage discrimination affected stability at certain parasitoid development times, but not at others. The relative lengths of parasitoid and host development times also influenced the tendency of the host population to show discrete or overlapping generations.     

The effect of egg load on readiness to accept a low-quality host plant is weak and age dependent in a geometrid moth

Abstract.  1. Most empirical and theoretical studies treat egg load as a major determinant of insect oviposition behaviour, and predict a positive link between egg load and readiness to oviposit.
2. In the present study, the correlation between egg load and readiness to lay eggs on a low-ranked host was examined in wild-caught moths, Scotopteryx chenopodiata L. (Lepidoptera: Geometridae).
3. No overall effect of egg load on oviposition could be detected. However, there was a significant interaction between egg load and wing wear: in only one wing-wear class, out of four, high egg load tended to promote oviposition, while in the three other classes the effect was insignificant.
4. The results suggest that the effect of egg load on oviposition may depend on the age of the ovipositing insect.     

The effects of host plant phenology on the demography and population dynamics of the leaf-mining moth, Cameraria hamadryadella (Lepidoptera: Gracillariidae)

Abstract.

• 1 We examined the effects of variation in the timing of spring leaf production and autumn leaf fall on the survival, mortality and abundance of Cameraria hamadryadella on Quercus alba and Q.macrocarpa.
• 2 We monitored and manipulated the timing of foliation on field and potted Q.alba trees and observed the abundance of C.hamadryadella on those trees. We also monitored and manipulated the timing of leaf fall on Q.alba and Q.macrocarpa trees in the field and observed its effects on survival, mortality and abundance of C.hamadryadella.
• 3 Variation in the timing of spring leaf production has no effect on C. hamudryadella abundance. However, a warm winter and spring in 1991 led to accelerated development and the imposition of a facultative third generation in one out of ten years of observation.
• 4 In 1989, leaves fell relatively early and leaf fall in the autumn accounted for more than 50% of the mortality of C.hamudryadella. in 1990 and 1991 leaves fell relatively late and leaf fall induced mortality was substantially reduced and overwinter survival was markedly increased.
• 5 The abundance of C.hamadryadella remained constant in the spring and summer of 1990 following the previous autumn's relatively early leaf fall, but increased by 10-fold in the spring of 1991 following the relatively late leaf fall of autumn 1990. The abundance of C.hamadryadella also increased 4-fold between the summer of 1991 and the spring of 1992 after another autumn of relatively late leaf fall. We attribute these increases in abundance in part to reduced mortality because of later leaf fall.
• 6 Variation in the timing of autumn leaf fall may be responsible for initiating outbreaks of C.hamadryadella.

     

Host-feeding and oviposition by parasitoids on hosts of different fitness value: Influences of egg load and encounter rate

A dynamic optimization model is presented for the decision to host feed or oviposit on hosts by female parasitoids. Optimal host utilization decisions are compared between two host types with different fitness payoffs for oviposition. The model predicts that hosts of higher fitness value should always be used for oviposition unless the egg load approaches zero. This general prediction is not influenced by levels of host availability or metabolic reserves, the age of the parasitoid, or the magnitude of the fitness difference. An egg-load threshold is predicted above which lower value hosts should be used for oviposition and below which they are used for host feeding. The position of this egg-load threshold is higher when the difference in fitness between host types is larger. The threshold is also higher when overall host availability is high or metabolic reserves for the production of new eggs are low. The threshold for oviposition on low-value hosts decreases to zero near the end of the parasitoid’s life. Under conditions where high-value hosts are rarely encountered compared to low-value hosts, the model predicts that lower value hosts should be accepted for oviposition at a lower egg-load threshold.     

Effect of host deprivation on egg quality,egg load,and oviposition in a solitary parasitoid,Chetogena edwardsii (Diptera: Tachinidae)

Solitary parasitoids are limited to laying one egg per host because larvae compete within hosts. If host encounter rate is low, females should not increase the number of eggs/host in response. The tachinid fly, Chetogena edwardsii,was used to evaluate the effect of host deprivation on egg accumulation, oviposition behavior, and egg quality in a solitary parasitoid. Females deprived of hosts for 2– 7 days accumulate about 1 day's supply of eggs. Egg output of deprived females once hosts are restored does not differ from that of control females. Deprived females retain one egg in the uterus where it undergoes embryogenesis. Maggots emerging from retained eggs are more likely to survive in hosts molting in 40 h or less after receipt of an egg than are maggots emerging from eggs fertilized shortly before oviposition. Egg retention is a consequence of host deprivation that permits females to broaden the range of hosts they can exploit to include soon-to-molt hosts and possibly multiply parasitized hosts.     

Influence of host deprivation on egg load and oviposition behaviour of Brachymeria intermedia, a parasitoid of gypsy moth

Abstract. Mated female Brachymeria intermedia (Hymenoptera: Chalcididae) deprived, since emergence, from pupae of their host Lymantria dispar (Lepidoptera: Lymantriidae), accumulated eggs but had a very low rate of hostacceptance. Parasitoids that were host-deprived after encountering pupae early in life also accumulated eggs, but maintained a high acceptance rate. Thus early exposure to hosts promoted active reproductive behaviour. Total egg production depended on the total number of pupae encountered, indicating that B.intermedia adjust their egg production to host availability. Hence, in B. intermedia both the physiological state of the parasitoid (age and egg load) and the informational state (in this case host-availability and experience) interact to shape oviposition behaviour.     

The oviposition behavior ofAphidius ervi andMonoctonus paulensis (Hymenoptera: Aphididae) encountering different host species (Homoptera: Aphididae) in sequential patches

Two-day-old mated females ofAphidius ervi Haliday andMonoctonus paulensis (Ashmead) were each provided with two sequential host patches. Patches were comprised of plastic petri dishes containing either 15 pea aphids,Acyrthosiphum pisum (Harris), or 15 alfalfa aphids,Macrosiphum creelii Davis. Both wasp species parasitized more hosts in patches containing pea aphids than in those containing alfalfa aphids, regardless of sequence. Females ofA. ervi also laid more eggs per aphid in patches containing pea aphids than in patches containing alfalfa aphids. When both patches contained alfalfa aphids,M. paulensis females parsitized more aphids in the second patch than in the first. Fewer alfalfa aphids were parasitized in the second patch when the first patch contained pea aphids, and fewer eggs were laid per alfalfa aphid. Parasitoid females of both species exhibited consistently higher rates of oviposition into their preferred host species and adjusted their reproductive allocation to hosts and host patches as a function of their experience in previous patches.     

Importance of host oviposition pattern and plant size for the selection of Trichogramma strains to control the diamondback moth

The aim of this work was to select a candidate strain of Trichogrammatidae (Hymenoptera: Chalcidoidea) to control the diamondback moth, Plutella xylostella L. (Lepidoptera: Plutellidae), an important pest of cabbage in Europe. The parasitic efficiency of Trichogramma chilonis Ishii from Japan and Réunion Island, Trichogramma evanescens Westwood from Egypt, Trichogramma ostriniae Pang & Chen from Japan, and Trichogramma semblidis (Aurivillius) from France was studied with sentinel eggs in greenhouse‐grown cauliflower. The percentage of parasitized eggs was measured 1 m from the release point and 3 days after release. Two factors were studied: host‐egg density (three eggs per stem vs. 50 eggs per stem) and plant size (50–100 cm high plants vs. 100–150 cm high plants). Trichogramma evanescens from Egypt achieved the best parasitism. Parasitism efficiency of T. ostriniae from Japan and T. chilonis from Réunion Island was not influenced by egg density. In contrast, parasitism efficiencies of the three other strains were higher at the lower egg density than at the higher egg density. Parasitism efficiency was lower when the plants were bigger, with the exception of T. chilonis from Réunion Island, for which the level of parasitism was not affected by plant size. Thus, T. chilonis from Réunion Island appeared to be the most suitable Trichogramma strain for use in controlling P. xylostella, regardless of the growth stage of the crop. However, T. ostriniae from Japan may also be more suitable for use on cauliflower crops at the early stage of crop development. If potential problems related to the introduction of exotic species are taken into consideration, it is possible to consider the native T. evanescens.     

Phenotypic divergence in reproductive traits of a moth population experiencing a phenological shift

Allochrony that is reproductive isolation by time may further lead to divergence of reproductive adaptive traits in response to different environmental pressures over time. A unique “summer” population of the pine processionary moth Thaumetopoea pityocampa, reproductively isolated from the typical winter populations by allochronic differentiation, is here analyzed. This allochronically shifted population reproduces in the spring and develops in the summer, whereas “winter” populations reproduce in the late summer and have winter larval development. Both summer and winter populations coexist in the same pine stands, yet they face different climatic pressures as their active stages are present in different seasons. The occurrence of significant differences between the reproductive traits of the summer population and the typical winter populations (either sympatric or allopatric) is thus hypothesized. Female fecundity, egg size, egg covering, and egg parasitism were analyzed showing that the egg load was lower and that egg size was higher in the summer population than in all the studied winter populations. The scales that cover the egg batches of T. pityocampa differed significantly between populations in shape and color, resulting in a looser and darker covering in the summer population. The single specialist egg parasitoid species of this moth was almost missing in the summer population, and the overall parasitism rates were lower than in the winter population. Results suggest the occurrence of phenotypic differentiation between the summer population and the typical T. pityocampa winter populations for the life‐history traits studied. This work provides an insight into how ecological divergence may follow the process of allochronic reproductive isolation.     

The distribution of eggs per host in a herbivorous insect--intersection of oviposition, dispersal and population dynamics

1. The dynamics of parasitic organisms depend critically upon the frequency distribution of parasite individuals per host. However, the processes giving rise to this frequency distribution have rarely been modelled and tested for organisms with complex host selection behaviour. 2. In this study Microrhopala vittata, a chrysomelid beetle, was used to investigate how oviposition behaviour, movement and density of host plants interact in shaping the frequency distribution of egg clusters per host in the field. 3. Enclosures were stocked with two different host species and different beetle densities and various stochastic process models were fitted to egg cluster count data obtained from these enclosures. The different models were derived considering different scenarios, in particular whether or not plant density limits oviposition rate, whether or not ovipositing females actively seek out the most attractive plant within their perception radius and whether a female's oviposition rate is determined by plant intrinsic factors, the plant's egg cluster load or the surrounding beetle density. 4. The model parameters fitted to cage data were used to describe the frequency distribution of egg cluster counts obtained in a release experiment in the field. A total of 220 beetle pairs were released at five locations in a field where this beetle was not observed previously. Each release point was at a border between the two host species. 5. One model predicted for the preferred host species the egg cluster count frequencies in the field from parameters estimated in the cages. This model assumed that egg clusters present on a plant increased subsequent oviposition on this plant. All other models could not describe the distribution of egg cluster counts for either of the two host species. 6. The results suggest that females seek out attractive hosts actively and the attractiveness of a plant increases with its egg cluster load. This behaviour creates a frequency distribution of egg clusters per host that depends only on beetle density but not on plant density. This conclusion has important implications for modelling insect-plant interactions.     

Identification of chemical oviposition stimulants for the diamondback moth, Plutella xylostella, present in three species of Brassicaceae

Plant chemicals in three cruciferous crop species, Brassica napus L., B. juncea (L.) Czerniak, and Sinapis alba L., that stimulate oviposition in the diamondback moth, Plutella xylostella (L.) (Lepidoptera: Plutellidae) were investigated in laboratory bioassays. Aerial portions of 4- to 6-week-old plants were extracted and fractionated using ion-exchange liquid chromatography. The oviposition stimulants were identified as glucosinolates, which are found in all Brassicaceae species. Activity of extracts was largely eliminated by treatment with myrosinase or sulphatase, enzymes which degrade glucosinolates. Reference standards of the same glucosinolates and in the same concentrations as in the extracts were equally stimulatory. A test with eight different glucosinolates demonstrated that the moths do not discriminate between glucosinolates with different side-chain structures. However, in tests using allylglucosinolate the oviposition response was dose-dependent. One of the species tested, S. alba, contained a possible oviposition deterrent.

---

Résumé Les produits chimiques trouvés dans trois espèces de crucifères cultivées, Brassica napus L., B. juncea (L.) Czerniak, et Sinapis alba L., qui stimulent l'oviposition chez la teigne des crucifères, Plutella xylostella (L.) (Lepidoptera: Plutellidae) ont été examinés. La partie aérienne des plants agés de 4 à 6 semaines a été extraite avec le methanol bouillant à 80%, le methanol pur, et l'éther. La concentration par filtration sur célite a donné comme resultat un extrait aqueux. Le materiel restant sur la célite a été dissout avec hexane pour donner un extrait lipophillique. L'extrait aqueux a été fractionné à l'aide de la chromatographie liquide sur échangeur d'ion pour donner trois fractions: neutre, cationique, et anionique. Les extraits et les fractions ont été ajustés à 1 g poids frais de tissu de plant par ml, appliqués sur du papier filtre, et exposés aux papillons femelles dans les essais de choix d'oviposition. L'oviposition a été stimulée fortement en présence de l'extrait aqueux, la fraction anionique, et quelque sous-fractions anioniques. Plus tard, il a été déterminé que ces derniers contenaient des glucosinolates.Chez les trois espèces de crucifères, les stimulants d'oviposition ont été identifiés comme étant des glucosinolates, que l'on retrouve dans toutes les espèces de crucifères. L'activité des extraits a été éliminée en grande partie par traitement avec myrosinase ou sulphatase, des enzymes qui dégradent spécifiquement les glucosinolates. Des standards de references des mêmes glucosinolates et aux même concentrations que dans les extraits ont eu également un effet stimulant. Un essai avec huit glucosinolates differentes à une concentration de 50 g/ml appliquées sur du papier filter à 3.2 g/cm² a démontré que les papillons ne discriminent pas entre les glucosinolates possédant des chaines secondaires differentes. Par contre, dans les essais utilisant l'allylglucosinolate, la réponse d'oviposition a été dépendente de la dose. S. alba a semblé contenir un inhibiteur de l'oviposition, qui est retrouvé dans l'extrait aqueux mais non pas dans la fraction anionique.

     

Induction of plant responses to oviposition and feeding by herbivorous arthropods: a comparison

Plants may respond both to feeding and oviposition by herbivorous insects. While responses of plants to feeding damage by herbivores have been studied intensively during the past decades, only a few, but growing number of studies consider the reactions of plants towards egg deposition by herbivorous insects. Plants showing defensive response to oviposition by herbivores do not `wait' until being damaged by feeding, but may instead react towards one of the initial steps of herbivore attack, the egg deposition. Direct plant defensive responses to feeding act directly against the feeding stages of the herbivores. However, a plant may also show direct defensive responses to egg deposition by (a) formation of neoplasms, (b) formation of necrotic tissue (= hypersensitive response), and (c) production of oviposition deterrents. All these plant reactions have directly negative effects on the eggs, hatching larvae, or on the ovipositing females. Indirect plant defensive responses to feeding result in the emission of volatiles (= synomones) that attract predators or parasitoids of the feeding stages. A few recent studies have shown that plants are able to emit volatiles also in response to egg deposition and that these volatiles attract egg parasitoids. Studies on the mechanisms of induction of synomones by egg deposition show several parallels to the mechanisms of induction of plant responses by feeding damage. When considering induced plant defence against herbivores from an evolutionary point of view, the question arises whether herbivores evolved the ability to circumvent or even to exploit the plant's defensive responses. The reactions of herbivores to oviposition induced plant responses are compared with their reactions to feeding induced plant responses.     

Control of reproduction and a survival cost to mating in female Bicyclus anynana butterflies

Abstract.  1. Given that sexual conflict is all pervasive, investigating potential costs to mating and the control of female reproduction are important issues.
2. Here, female reproductive output and longevity are investigated in relation to mating status (virgin vs. once-mated females) and host-plant availability in the tropical butterfly Bicyclus anynana .
3. Both factors significantly affected realised fecundity in a manner demonstrating that female reproduction is not entirely under control of the female herself, but is rather subject to additive contributions of the female, her male partner, and intrinsic physiological processes. As evident from the deposition of significant egg numbers by virgin, host-deprived females, B. anynana is effectively unable to completely inhibit oogenesis.
4. Mated females suffered a reduction in adult life span, which cannot be explained as a side-effect of variation in egg size, lifetime, or early fecundity.
5. Such detrimental effects of mating per se are indicative of the cooperation–conflict balance between sexes being shifted towards conflict in B. anynana .     

Changes in egg colour, egg weight and oviposition rate with the number of eggs laid by wild females of the small heath butterfly, Coenonympha pamphilus

ABSTRACT. 1. The colour of eggs laid by individual Coenonympha pamphilus (L.) (Lep., Satyridae) females changed over their lifespan. The first laid eggs were green, but after having laid about 100 eggs, females laid only yellow eggs.
2. By following females in the field, or by capturing wild females, and noting the colour of their eggs, it was established that younger females showed higher oviposition rates and laid heavier eggs than older females.
3. Hence, wild C.pamphilus females are unable to sustain a constant egg production by extracting amino acids or any other nutrients from nectar. We hypothesize that this is a general phenomenon in nectar feeding butterflies.
4. The egg colour did not seem to match the colour of the substrate on which the female chose to deposit the egg.