Host‐associated divergence in the activity of digestive enzymes in two populations of the gypsy moth Lymantria dispar (Lepidoptera: Erebidae)

The gypsy moth is a generalist insect pest with an extremely wide host range. Adaptive responses of digestive enzymes are important for the successful utilization of plant hosts that differ in the contents and ratios of constituent nutrients and allelochemicals. In the present study, we examined the responses of α‐amylase, trypsin, and leucine aminopeptidase to two tree hosts (suitable oak, Quercus cerris, and unsuitable locust tree, Robinia pseudoacacia) in the fourth, fifth, and sixth instars of gypsy moth larvae originating from oak and locust tree forest populations (hereafter assigned as Quercus and Robinia populations, respectively). Gypsy moths from the Robinia forest had been adapting to this unsuitable host for more than 40 generations. To test for population‐level host plant specialization, we applied a two‐population × two‐host experimental design. We compared the levels, developmental patterns, and plasticities of the activities of enzymes. The locust tree diet increased enzyme activity in the fourth instar and reduced activity in advanced instars of the Quercus larvae in comparison to the oak diet. These larvae also exhibited opposite developmental trajectories on the two hosts, i.e. activity increased on the oak diet and decreased on the locust tree diet with the progress of instar. Larvae of the Robinia population were characterized by reduced plasticity of enzyme activity and its developmental trajectories. In addition, elevated trypsin activity in response to an unsuitable host was observed in all instar larvae of the Robinia population, which demonstrated that Robinia larvae had an improved digestive performance than did Quercus larvae.     

Estimating the susceptibility of tree species to attack by the gypsy moth, Lymantria dispar

ABSTRACT.

• 1 Numbers of gypsy moth larvae feeding on each of 922 randomly sampled trees in a Quercus—Acer—Fraxinus forest in southwestern Quebec, Canada were counted in 1979 and in 1980 to quantify the larval feeding preferences as observed in the field for eighteen deciduous and one coniferous tree species at the northern range limit of the gypsy moth.
• 2 Both the diameter at breast height (dbh) and the estimated foliage biomass of the sampled trees were used to calculate the relative proportions of foliage represented by each of the nineteen tree species in the forest canopy. With these data on availability and utilization of the tree species by the gypsy moth larvae an Ivlev-type electivity index was used to quantify the larval feeding preferences. These preferences observed in the field define the susceptibility of a tree species to attack by the gypsy moth.
• 3 The feeding preferences calculated using estimated foliage biomass were comparable to the simpler calculation based on dbh (Spearman's rho = 0.79; P= 0.0001). The dbh-based feeding preferences remained almost unchanged in 1979 and 1980 (Spearman's rho = 0.83; P= 0.0001).
• 4 The composite 1979—80, dbh-based feeding preferences show Quercus rubra, Populus grandidentata, Ostrya virginiana, Amelanchier spp. and Acer saccharum were preferentially attacked by gypsy moth. Prunus serotina, Betula lutea, Acer rubrum, A. pensylvanicum, Fraxinus americana, Ulmus rubra, P. pensylvanicum and B. papyrifera were avoided. All nineteen tree species were, however, utilized to at least some degree by gypsy moth larvae.
• 5 These results quantitatively affirm and clarify earlier reports of gypsy moth feeding preferences in North America and Eurasia. The advantages and limitations of using an electivity index to estimate the susceptibility of different tree species to attack by folivores like the gypsy moth are discussed.

     

Suitability of Eucalyptus and Corymbia for Mnesampela privata (Guenée) (Lepidoptera: Geometridae) larvae

Abstract 1 Mnesampela privata (Guenée) has a host list of 40 Eucalyptus and at least one Corymbia species. Larval survival and performance was studied on 19 species to investigate how certain leaf traits influence the suitability of different species. 2 After 7 days, survival on Eucalyptus aggregata and Eucalyptus camphora is greater than 70% even though the toughness of leaves is 0.15–0.19 mg/mm². However, after the same time, survival on genotypes of Eucalyptus melliodora and Eucalyptus sideroxylon was less than 60%, or even 0%, even though the toughness of some leaves was as low as 0.11 mg/mm². An unmeasured allelochemical, rather than toughness, may reduce survival on these species. 3 Dry weights of first‐instar larvae were negatively correlated with leaf toughness for 13 of the species studied. Species that produced the heaviest first‐instar larvae were not the same hosts that produced the heaviest second‐instar larvae. 4 Dry weights of female pupae were negatively correlated with total oil content for five of the species studied. 5 Larvae exhibit age‐related changes in feeding behaviour. Neonates skeletonize leaves (avoid leaf veins and oil glands) and post‐third‐instar larvae ingest whole leaf fragments (consume small leaf veins and oil glands). These findings suggest that neonates are sensitive to high leaf toughness and non‐oil plant secondary metabolites whereas older larvae are less sensitive to high leaf toughness and are likely to become larger adults on hosts with lower oil contents.     

Growth responses of gypsy moth larvae to elevated CO2: the influence of methods of insect rearing

Abstract The effects of elevated CO₂ on foliar chemistry of two tree species (Populus pseudo‐simonii Kitag. and Betula platyphylla) and on growth of gypsy moth (Lymantria dispar L.) larvae were examined. Furthermore, we focused on the comparison of results on the growth responses of larvae obtained from two methods of insect rearing, the no‐choice feeding trial performed in the laboratory or in situ in open‐top chambers. On the whole, both primary and secondary metabolites in the leaves of the two tree species were significantly affected by main effects of time (sampling date), CO₂ and species. Elevated CO₂ significantly increased the C : N ratio and concentrations of the soluble sugar, starch, total nonstructural carbohydrates, total phenolics and condensed tannins, but significantly decreased the concentration of nitrogen. Higher contents of total phenolics and condensed tannins were detected in the frass of larvae reared in elevated CO₂ treatments. Overall, the growth of gypsy moth larvae were significantly inhibited by elevated CO₂ and CO₂‐induced changes in leaf quality. Our study did not indicate the two methods of insect rearing could influence the direction of effects of elevated CO₂ on the growth of individual insects; however, the magnitude of negative effects of elevated CO₂ on larval growth did differ between the two insect rearing methods, and it seems that the response magnitude was also mediated by larval age and host plant species.     

Interspecific competition between the braconid endoparasitoids Glyptapanteles porthetriae and Glyptapanteles liparidis in Lymantria dispar larvae

The effect of interspecific competition between the solitary endoparasitoid Glyptapanteles porthetriae Muesebeck (Hymenoptera: Braconidae) and the gregarious Glyptapanteles liparidis Bouché (Hymenoptera: Braconidae), was investigated in larvae of Lymantria dispar L. (Lepidoptera: Lymantriidae). Host larvae were parasitized by both wasp species simultaneously in premolt to the 2^nd or the 3^rd host instar or in an additional approach with a 4‐day delay in parasitization by the second wasp species. Host acceptance experiments revealed that both wasp species do not discriminate between unparasitized host larvae and larvae parasitized previously by the same or the other species. In more than 90% female wasps parasitized the larva they encountered first. During the period of endoparasitic development, larvae of the competing parasitoid species never attacked the egg stage of the other species. When host larvae were parasitized simultaneously by both wasp species, the rate of successful development of both species depended on the age of the host larva at the time of its parasitization; G. liparidis emerged successfully from 44% of host larvae parasitized during the premolt to 2^nd instar, G. porthetriae from 28%, and in 20% of the hosts both parasitoid species were able to develop in one gypsy moth larva. However, when host larvae were parasitized simultaneously during premolt to the 3^rd instar, G. liparidis was successful in 90% of the hosts, compared to 8% from which only G. porthetriae emerged. In the experiments with delayed oviposition, generally the species that oviposited first succeeded in completing its larval development. Larvae of the species ovipositing with four days delay were frequently attacked and killed by larvae of the first parasitizing species or suffered reduced growth. As the secondary parasitoid species, G. porthetriae‐larvae were never able to complete their development, whereas G. liparidis developed successfully in at least 12,5% of the multiparasitized host larvae. Thus, multiparasitism of gypsy moth larvae by both Glyptapanteles species corresponds to the contest type; however, G. porthetriae is only able to develop successfully as the primary parasitoid of young host larvae.     

Different susceptibility of indigenous populations of Lymantria dispar to the exotic entomopathogen Entomophaga maimaiga

The recovery of the host‐specific entomopathogen Entomophaga maimaiga is still limited to certain world areas, although it is recently spreading to Eastern Europe. This study evaluated the effectiveness and fitness of an E. maimaiga isolate from Balkans against Lymantria dispar populations collected along the Italian peninsula and main islands, where the fungus has never been reported. As a result of different bioassays, the pathogenicity against gypsy moth larvae was generally confirmed, although significant differences among insects feeding upon diverse forest plant species were observed. The lack of significant susceptibility of other lepidopteran species from the same areas is also reported.     

Interspecific competition between the braconid endoparasitoids Glyptapanteles porthetriae and Glyptapanteles liparidis in Lymantria dispar larvae

The effect of interspecific competition between the solitary endoparasitoid Glyptapanteles porthetriae Muesebeck (Hymenoptera: Braconidae) and the gregarious Glyptapanteles liparidis Bouché (Hymenoptera: Braconidae), was investigated in larvae of Lymantria dispar L. (Lepidoptera: Lymantriidae). Host larvae were parasitized by both wasp species simultaneously in premolt to the 2^nd or the 3^rd host instar or in an additional approach with a 4-day delay in parasitization by the second wasp species. Host acceptance experiments revealed that both wasp species do not discriminate between unparasitized host larvae and larvae parasitized previously by the same or the other species. In more than 90% female wasps parasitized the larva they encountered first. During the period of endoparasitic development, larvae of the competing parasitoid species never attacked the egg stage of the other species. When host larvae were parasitized simultaneously by both wasp species, the rate of successful development of both species depended on the age of the host larva at the time of its parasitization; G. liparidis emerged successfully from 44% of host larvae parasitized during the premolt to 2^nd instar, G. porthetriae from 28%, and in 20% of the hosts both parasitoid species were able to develop in one gypsy moth larva. However, when host larvae were parasitized simultaneously during premolt to the 3^rd instar, G. liparidis was successful in 90% of the hosts, compared to 8% from which only G. porthetriae emerged. In the experiments with delayed oviposition, generally the species that oviposited first succeeded in completing its larval development. Larvae of the species ovipositing with four days delay were frequently attacked and killed by larvae of the first parasitizing species or suffered reduced growth. As the secondary parasitoid species, G. porthetriae-larvae were never able to complete their development, whereas G. liparidis developed successfully in at least 12,5% of the multiparasitized host larvae. Thus, multiparasitism of gypsy moth larvae by both Glyptapanteles species corresponds to the contest type; however, G. porthetriae is only able to develop successfully as the primary parasitoid of young host larvae.     

Potential risk of accidental introduction of Asian gypsy moth (Lymantria dispar) to Australasia: effects of climatic conditions and suitability of native plants

1 The potential risk of the establishment of the Asian strain of the gypsy moth (AGM) (Lymantria dispar) in New Zealand and Australia (Australasia) was assessed from a study of the insect's host range and potential distribution. In New Zealand, viable eggs of AGM have been continuously intercepted on cargo from Asia, and therefore there is a high probability of accidental introductions of AGM to Australasia. 2 We predicted potential distribution ranges of AGM based on climatic conditions. Asian gypsy moth is predicted to be able to persist in N and SE New Zealand and SE and SW Australia. 3 Using three populations of AGM and 59 species (seven families) of plant (55 from Australasia and four from elsewhere), we also conducted laboratory trials to examine the ability of AGM larvae to complete development on native plants from Australasia. Asian gypsy moth was able to complete development on 26 out of the 55 native species tested. Furthermore, larval performance on at least five species of Australian native plant was as good as on AGM's preferred host species (Quercus pubescens and Q. robur). 4 Larval performance of AGM was poor on all but one species of New Zealand native tree species (Nothofagus solandri), and therefore the risk of establishment in the indigenous forests of New Zealand is considered to be low. 5 Given the suitability of some Australian plants and the climatic suitability for the establishment of AGM, this insect should be treated as a serious quarantine threat and managed accordingly, particularly in Australia.     

Larval performance of the oriental fruit moth across fruits from primary and secondary hosts

A common characteristic of many invasive herbivorous insects is their ability to utilize a broad range of host plants. By using various hosts in phenological succession, multivoltine herbivores may increase the number of successful annual generations, at the same time as potentially increasing their overall fitness. To achieve such success, herbivores must be able to develop efficiently on the nutritional resources offered by their hosts. The oriental fruit moth Cydia (= Grapholita) molesta (Busck) (Lepidoptera: Tortricidae) is one of the most damaging invasive insect species. Peach (Prunus persica) is its primary host, whereas the pome fruits apple (Malus × domestica Borkh) and pear (Pyrus communis) are considered as secondary hosts. In many parts of its geographical range, including southern Europe, populations of the moth switch from peach to apple or pear orchards during the growing season. The present study tests whether this temporal switch is supported by the physiological capability of the larvae with respect to developing efficiently on fruits of these taxonomically‐related host plants. Larvae are reared on peach, apple or pear fruits; several life‐history traits are measured; and correlations between the traits are calculated. The results obtained show that larvae do not have the same physiological capability with respect to using apple or pear fruits as hosts compared with using peach fruit. Pear fruit in particular is a sub‐optimal diet. These findings suggest that, in the case of continuous geographical expansion, concomitantly with global warming, apple orchards might support oriental fruit moth populations better than pear orchards, and that the switch onto novel hosts might be accompanied by restricted population growth.     

Host recognition by the gypsy moth [Lymantria dispar] [Lep.: Lymantriidae] hyperparasite,Eurytoma appendigaster [Hym.: Eurytomidae] of cocoons ofCotesia melanoscela [Hym.: Braconidae]

Artificial and modified natural hosts were exposed to females of the gypsy moth [Lymantria dispar (L.)] hyperparasite,Eurytoma appendigaster (Swederus), to investigate its host recognition behavior on the primary host, which are cocooned larvae of the gypsy moth parasite,Cotesia melanoscela (Ratzeburg). Material(s) which caused drilling behavior by the hyperparasite on host cocoons were extracted with both polar and non-polar solvents. However, cocoons washed with large volumes of solvent still caused substantial drilling activities by females, suggesting that additional cues may be important. Results suggest that host recognition in this hyperparasite involves a variety of host characteristics.      

Nutritional indices in the gypsy moth (Lymantria dispar (L.)) under field conditions and host switching situations

A large proportion of gypsy moths (Lymantria dispar (L.)) are likely to experience multiple species diets in the field due to natural wandering and host switching which occurs with these insects. Nutritional indices in fourth and fifth instar gypsy moth larvae were studied in the field for insects that were switched to a second host species when they were fourth instars. The tree species used as hosts were northern pin oak (Quercus ellipsoidalis E. J. Hill), white oak (Q. alba L.), big-tooth aspen (Populus grandidentata Michx.), and trembling aspen (P. tremuloides Michx.). Conclusions of this study include: 1) Insects which fed before the host switch on northern pin oak performed better after the host switch than did insects with other types of early dietary experience. While the northern pin oak-started insects had very low relative food consumption rates on their second host species immediately after the switch, one instar later they had the highest ranked consumption rates. During both instars they had the second highest efficiencies of converting ingested and digested food to body mass. High food consumption rates and relatively high efficiency of food conversion helped these insects to obtain the highest ranked mean relative growth rates in the fifth instar compared to the relative growth rates obtained by insects from any of the other first host species. 2) Among the four host species examined, a second host of trembling aspen was most advantageous for the insects. Feeding on this species after the switch led to higher larval weights and higher relative growth rates for insects than did any of the other second host species. The insects on trembling aspen attained excellent growth despite only mediocre to low food conversion efficiencies. The low efficiencies were offset by high relative food consumption rates. 3) Low food consumption rates often tend to be paired with high efficiency of conversion and vice versa. 4) There is no discernable tendency for the first plant species eaten to cause long-term inductions which affect the ability of gypsy moths to utilize subsequent host plants. Insects did not tend to consume more, grow faster, or be more efficient if their second host plant was either the same as their rearing plant or congeneric to it. Methods are delineated which allow values of nutritional indices to be obtained for insects on intact host plants under field conditions. These methods are useful for the purpose of answering questions about the relative effects that different diet treatments have on insect response.     

What physiological processes permit insects to eat Eucalyptus leaves?

Many species of insects eat Eucalyptus foliage despite its relatively low nutritional value and the many plant secondary metabolites (PSMs) present, for example, terpenes, phenols and formylated phloroglucinols (FPGs). Formylated phloroglucinols are a new class of PSMs that act as antifeedants for possums and koalas. What physiological processes are present that permit insects to eat eucalypt foliage and how do PSMs influence insect feeding or digestion? Some trees seem to be repeatedly infested with eucalypt‐feeding insects, possibly as a result of previous chemosensory cues remaining from parental selection of a plant. Avoidance or storage of PSMs permit jarrah leafminers (Perthida glyphopa) and sawflies (Perga sp.) to consume eucalypt foliage without dealing with the majority of these compounds. Some PSMs can be metabolized by polysubstrate membrane oxidases as found in caterpillars or sawflies that feed on eucalypts. High midgut pH may be advantageous for nutrient extraction and PSM metabolism, and midgut pH ranges between 8.5 and 8.9 for caterpillars of Hyalarcta huebneri. Plant secondary metabolites may not be absorbed as a result of the combined presence of the peritrophic matrix and endogenous surfactants. Excretion of PSMs can be as metabolites or intact compounds. Both putative metabolites and sideroxylonal‐A, an FPG, are present in the faeces of larvae of the case moth, H. huebneri. The presence of sideroxylonal‐A in the food had an effect on the presence of 5‐hydroxytryptamine (5HT) in the central nervous system of caterpillars, as larvae fed leaves with a high concentration of sideroxylonal‐A had relatively more 5HT in the brain and central nervous system ganglia than larvae fed leaves containing a low concentration. Further work is necessary to clarify how PSMs are handled by eucalypt‐feeding insects and what effect FPGs have on feeding and digestion.     

Rapid induced resistance of silver birch affects both innate immunity and performance of gypsy moths: the role of plant chemical defenses

In this study we tested the effects of rapid induced resistance of the silver birch, Betula pendula, on the performance and immune defense of the gypsy moth, Lymantria dispar. We also measured the effects of defoliation on the concentrations of plant secondary metabolites, particularly on phenolics and terpenoids. It was found that severe natural defoliation (by moth larvae) of silver birch led to an increase in lipophilic flavonoids on the leaf surface. The concentration of some simple phenolics and monoterpenes (linalool and geraniol) also increased, while that of several glycosides of quercetin decreased. The female pupal weights and survival rates of moths decreased, and larval development time increased, when the insects fed on defoliated trees. However, the feeding of caterpillars with the leaves of defoliated trees led to an increase in lysozyme-like activity in their hemolymph, with an increase in their ability to encapsulate potential parasites. Our data show that the silver birch deploys a rapid chemical defense against gypsy moth larvae. We suggest that lipophilic flavonoids are important compounds in the direct silver birch defense against L. dispar caterpillars. The increased strength of immune defense of insects exposed to trees that had deployed a rapid induced resistance may be an adaptation of the herbivores to resist the rising density of parasites when host population density is high.     

Host population structure affects field sex ratios of the heteronomous hyperparasitoid, Coccophagus atratus

Abstract.

• 1 Second instar Filippia gemina de Lotto scale insects are the preferred hosts of female Coccophugus atrutus Compere larvae. These scale insects were found on their host plants, Chrysanthemoides monilifera Norlindh and Cliffortia strobilifera Mettenius, only at certain times during a 1 year sampling programme.
• 2 Late larval instars and prepupae of C.atratus, and a Metaphycus species, are the preferred hosts of male C.atratus larvae. These hosts, although they occurred on the same host plants as hosts for female C.atratus, were most numerous at different times during the sampling period.
• 3 The ratio of hosts suitable for C.atratus varied from a predominance of hosts suitable for females through to a predominance of hosts suitable for males. Sex ratios of adult C.atratus followed a similar trend but did not reflect, exactly, the ratio of available hosts. Differences in mortality between sexes and hyperparasitism may account for this anomaly.
• 4 Variable population sex ratios observed in C.atratus apparently result from the behaviour of individual females in which brood sex ratios are dependent on the relative availability of hosts for males and hosts for females. This behaviour, in turn, may result from variability in the host population structure but may also result from selection pressures operating at the time that heteronomous hyperparasitism evolved.

     

Responses of a native beetle to novel exotic plant species with varying invasion history

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Effects of host switching on gypsy moth (Lymantria dispar (L.)) under field conditions

Effects of various single and two species diets on the performance of gypsy moth (Lymantria dispar (L.)) were studied when this insect was reared from hatch to population on intact host trees in the field. The tree species used for this study were red oak (Quercus rubra L.), white oak (Q. alba L.), bigtooth aspen (Populus grandidentata Michaux), and trembling aspen (P. tremuloides Michaux). These are commonly available host trees in the Lake States region. The study spanned two years and was performed at two different field sites in central Michigan. Conclusions drawn from this study include: (1) Large differences in gypsy moth growth and survival can occur even among diet sequences composed of favorable host species. (2) Larvae that spent their first two weeks feeding on red oak performed better during this time period than larvae on all other host species in terms of mean weight, mean relative growth rate (RGR), and mean level of larval development, while larvae on a first host of bigtooth aspen were ranked lowest in terms of mean weight, RGR, and level of larval development. (3) Combination diets do not seem to be inherently better or worse than diets composed of only a single species; rather, insect performance was affected by the types of host species eaten and the time during larval development that these host species were consumed instead of whether larvae ate single species diets or mixed species diets. (4) In diets composed of two host species, measures of gypsy moth performance are affected to different extents in the latter part of the season by the two different hosts; larval weights and development rates show continued effects of the first host fed upon while RGRs, mortality, and pupal weights are affected strongly by the second host type eaten. (5) Of the diets investigated in this study, early feeding on red oak followed by later feeding on an aspen, particularly trembling aspen, is most beneficial to insects in terms of attaining high levels of performance throughout their lives.     

Comparisons of mountain pine beetle (Dendroctonus ponderosae Hopkins) reproduction within a novel and traditional host: effects of insect natal history,colonized host species and competitors

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Host range testing for risk assessment of a sexually dimorphic polyphagous invader,painted apple moth

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Rapid evolution and specialization following host colonization in a yucca moth

Host shifts followed by specialization can result in sympatric genetic differentiation, and may have fuelled the diversification of phytophagous insects. This study examines a recent colonization of a non‐native host by Prodoxus quinquepunctellus (Lepidoptera: Prodoxidae). Allozyme differentiation was detected among different host feeding populations, yet was nearly absent among similar host feeding populations in sympatry. Geographical patterns of allozyme variation showed a much higher level of population structure among populations feeding on the derived host. Conversely, mtDNA haplotype frequencies were nearly homogeneous in the derived populations compared to the ancestral populations, suggesting a bottleneck and/or rapid fixation of haplotypes following host colonization. Moth emergence coincided with host plant flowering, and phenological differences between host species translated into allochronic isolation between populations feeding on different hosts. Derived moth populations also differed significantly in three ovipositor characters from ancestral populations. These findings suggest rapid host‐specific genetic differentiation, and specialization of moth emergence time and ovipositor morphology following host colonization.     

Colonization, survival, and causes of mortality of Camerariu hamadryadella (Lepidoptera: Gracillariidae) on four species of host plants

Abstract.

• 1 The abundance, survival, and causes of mortality of Cameraria hamadryadella (Clemens) (Lepidoptera: Gracillariidae) were examined on four host plant species in Virginia, U.S.A. Quercus alba L. and Q.rubra L. are native within the geographic range of C.hamadryadella, and Q.robur L. and Q.benderi Baenitz are exotic. Q.robur is native to Europe, North Africa, and Asia and was probably introduced prior to 1850, and Q.benderi is of hybrid origin and introduced to cultivation before 1900. Q.alba and Q.robur are in the subgenus Lepidobalanus (white oaks), and Q.rubra and Q.benderi are in the subgenus Erythrobalanus (red oaks).
• 2 Larval mines of C. hamadryadella were abundant on both white oak species, including the exotic Q.robur, but were rare on host plants in the red oak subgenus. Un-hatched eggs of C.hamadryadella were not observed on red oaks. Other observations on host distribution indicate that C.hamadryadella is rarely found on red oaks. These observations are interpreted as circumstantial evidence that C. hamadryadella generally avoids ovipositing on red oaks.
• 3 Survival of C.hamadryadella to the adult stage was similar among all host species, but larvae tended to survive longer on hosts in the red oak subgenus. The observation of higher survival rates of early instar larvae on red oaks suggests that no nutritional or secondary chemical barrier reinforces the observed pattern of oviposition.
• 4 Significant differences in the distribution of the causes of mortality were detected between native and exotic host plant species. Larvae and pupae on native hosts were more likely to die because of predation, while those on exotic host plants were more likely to die because of parasitism and host feeding by adult female parasitoids. This pattern could arise because parasitoids prefer to forage on exotic host plants or because predators avoid foraging on exotic plants.
• 5 This study shows for C. hamadryadella that the only barriers to colonization and use of exotic hosts, in the white and red oak subgenera, are the presence of cues sufficient to stimulate oviposition and/or the absence of cues to deter oviposition. It also suggests that the presence of closely related native host plants in the region of introduction will increase the probability that exotic plants will be colonized by phytophagous insects.