Nucleopolyhedroviruses of forest and western tent caterpillars: cross-infectivity and evidence for activation of latent virus in high-density field populations

Abstract. 1. Cyclic population dynamics of forest caterpillars are often associated with epizootics of nucleopolyhedrovirus, but it is not known how these viruses persist between generations or through the fluctuations in host population density. 2. To explore the question of virus persistence at different phases of the population cycle, the nucleopolyhedroviruses of two species of tent caterpillar that co‐occur in British Columbia, Canada, Malacosoma californicum pluviale (western tent caterpillar) and Malacosoma disstria (forest tent caterpillar), were characterised. The cross‐infectivity of the viruses in these two host species was investigated to determine whether there might be a route for virus persistence via the alternative host species. Any virus produced in the cross‐infections was characterised to confirm true cross‐infection or to ascertain whether cross‐inoculation triggered latent virus persisting within the population. 3. The virus associated with forest tent caterpillars (MadiNPV) did not infect western tent caterpillars from low‐density populations, nor did it trigger a latent virus infection; however, inoculation of forest tent caterpillars from high‐density populations with virus from western tent caterpillars (McplNPV) resulted in viral infection, but without a dose–response relationship. 4. Analysis of DNA profiles of virus resulting from cross‐infection of the forest tent caterpillar with McplNPV, revealed that 88% of these infections were caused by MadiNPV rather than McplNPV; however the virus from all 44 infected individuals was identical and differed in DNA profile from the stock MadiNPV used for cross‐infection. This suggests strongly that forest tent caterpillars from high‐density field populations harbour a latent, persistent, or sublethal form of MadiNPV that was triggered by exposure to nucleopolyhedrovirus from the western tent caterpillar. 5. Virus was not activated in western tent caterpillars collected over 2 years of late population decline and the first year of population increase.     

Population fluctuations of the western tent caterpillar in southwestern British Columbia

Cyclic populations of western tent caterpillars fluctuate with a periodicity of 6–11 years in southwestern British Columbia, Canada. Typically, larval survival is high in early stages of the population increase, begins to decline midway through the increase phase, and is low through several generations of the population decline. Fecundity is generally high in increasing and in peak populations but is also reduced during the population decline. Poor survival and low fecundity for several generations cause the lag in recovery of populations that is necessary for cyclic dynamics. The dynamics of tent caterpillar populations vary among sites, which suggests a metapopulation structure; island populations in the rainshadow of Vancouver Island have more consistent cyclic dynamics than mainland populations in British Columbia. Sudden outbreaks of populations that last a single year suggest that dispersal from source to sink populations may occur late in the phase of population increase. Wellington earlier discussed qualitative variation among tent caterpillar individuals as an aspect of population fluctuations. The variation in caterpillar activity he observed was largely statistically nonsignificant. Recent observations show that the frequency of elongate tents as described by Wellington to characterize active caterpillars varies among populations but does not change in a consistent pattern with population density. The level of infection from nucleopolyhedrovirus (NPV) was high in some populations at peak density but was not associated with all population declines. Sublethal infection can reduce the fecundity of surviving moths, and there is a weak association between viral infection and egg mass size in field populations. The impact of weather in synchronizing or desynchronizing populations is a factor to be investigated further. Received: May 25, 1999 / Accepted: March 28, 2000     

Hierarchical spatial structure of genetically variable nucleopolyhedroviruses infecting cyclic populations of western tent caterpillars

The cyclic population dynamics of western tent caterpillars, Malacosoma californicum pluviale, are associated with epizootics of a nucleopolyhedrovirus, McplNPV. Given the dynamic fluctuations in host abundance and levels of viral infection, host resistance and virus virulence might be expected to change during different phases of the cycle. As a first step in determining if McplNPV virulence and population structure change with host density, we used restriction fragment length polymorphism (RFLP) analysis to examine the genetic diversity of McplNPV infecting western tent caterpillar populations at different spatial scales. Thirteen dominant genetic variants were identified in 39 virus isolates (individual larvae) collected from field populations during one year of low host density, and another distinct variant was discovered among nine additional isolates in two subsequent years of declining host density. The distribution of these genetic variants was not random and indicated that the McplNPV population was structured at several spatial levels. A high proportion of the variation could be explained by family grouping, which suggested that isolates collected within a family were more likely to be the same than isolates compared among populations. Additionally, virus variants from within populations (sites) were more likely to be the same than isolates collected from tent caterpillar populations on different islands. This may indicate that there is limited mixing of virus among tent caterpillar families and populations when host population density is low. Thus there is potential for the virus to become locally adapted to western tent caterpillar populations in different sites. However, no dominant genotype was observed at any site. Whether and how selection acts on the genetically diverse nucleopolyhedrovirus populations as host density changes will be investigated over the next cycle of tent caterpillar populations.     

Climate and outbreaks of the forest tent caterpillar

Some have suggested that the periodic outbreaks of the forest tent caterpillar. Malacosoma disstria are triggered by weather the temperature at the time of early larval feeding, and overwintering temperatures To assess the role of these factors, defoliation maps, compiled annually from flight surveys for the province of Ontario. were compared to similarly scaled temperature records An analysis of the year to year variation shows no relationship between the pattern of increases or declines in defoliation and either temperatures during early larval development, or overwintering temperatures Four periods of defoliation by forest tent caterpillars were observed in Ontario over 41 yr. but at individual sites extensive defoliation did not occur for each of the outbreak periods Defoliation was less severe in regions with low overwintering temperatures, but was not related to the average number of degree days m the early spring Outbreaks were most common in areas where deciduous forests were extensive, and the mean overwintering temperatures were above −40°C While these weather variables do not apparently explain the details of population dynamics of forest tent caterpillars, extreme weather conditions might synchronize populations     

The dynamics of forest tent caterpillar outbreaks across east‐central Canada

An analysis of forest tent caterpillar Malacosoma disstria defoliation records from Ontario and Quebec over the period 1938–2002 indicates that outbreaks recur periodically and somewhat synchronously among regions of the two provinces. Cluster analysis revealed that the most strongly periodic, large‐scale, synchronized fluctuations occurred within three regions: northwestern Ontario, eastern Ontario/western Quebec and southeastern Quebec. Defoliation in the vast surrounding hinterlands tended to be infrequent and sporadic, loosely tracking defoliation in the core outbreak regions. One small cluster in northeastern Ontario stood out as anomalous, as a result of an increasing trend in the duration of periodic defoliation episodes, marked by an unprecedented double‐wave of defoliation that persisted from 1992 to 1999. This is the precise area where, in the early 2000s, trembling aspen Populus tremuloides stems were mapped as being in a state of decline of unprecedented severity and extent. Our results suggest forest tent caterpillar has the potential to cause significant impacts on forest health and, hence, carbon budgets in east‐central Canada and that the forest tent caterpillar deserves more attention as a model system of forest insect disturbance ecology.     

Does tent caterpillar attack reduce the food quality of red alder foliage?

Summary We assayed the quality of red alder trees for western tent caterpillar growth and survival to test the hypothesis that caterpillar feeding stimulates plant defenses in both attacked and adjacent trees. Three years of high tent caterpillar density were necessary before deterioration in foliage quality occurred, and even then only foliage from trees which were almost completely defoliated in the current year reduced the growth of caterpillars. Both tent size and mean egg mass size increased after the second year of high density which indicates that good conditions still existed for tent caterpillars after 2 to 3 years of heavy feeding.Egg masses which were moved to areas where trees had not recently supported a high caterpillar population produced significantly smaller tents than endemic controls in 1982. Therefore the small tent and egg mass size of the high density population in 1982 was inherent to the insects rather than modified by food source. In 1983 the tents from introduced egg masses were as large as naturally occurring tents.If lightly attacked trees within areas of high caterpillar density are better defended against insect attack, this does not show up in their ability to support caterpillar growth and survival. We found no evidence to support the hypothesis that trees communicate insect attack and stimulate chemical defenses in adjacent trees. Reduced foliage quality seems to be a result of extensive insect damage rather than a defense against insect damage.     

Effects of variable phytochemistry and budbreak phenology on defoliation of aspen during a forest tent caterpillar outbreak

1 The present study assessed the relationship between clonally variable rates of defoliation in trembling aspen (Populus tremuloides Michx.) and two potential resistance traits: defensive chemistry and leaf phenology. 2 In 2001, coincident with a major outbreak of the forest tent caterpillar (Malacosoma disstria Hubner) in the northcentral U.S.A., we monitored defoliation rates, phytochemical composition, and foliar development in 30 clones of trembling aspen. Leaf chemistry was also assessed in re‐flushed leaves and 2 years post‐outbreak. 3 Early in the season, differences in defoliation among clones were substantial but, by mid‐June, all clones were completely defoliated. Leaf nitrogen, condensed tannins, and phenolic glycosides varied among clones but did not relate to defoliation levels. Budbreak phenology differed by 3 weeks among clones and clones that broke bud early or late relative to forest tent caterpillar eclosion experienced reduced rates of defoliation. 4 Defoliation led to increased tannins and slight decreases in phenolic glycoside concentrations in damaged leaf remnants, but to moderately decreased tannins and a six‐fold increase in phenolic glycosides in reflushed leaves. This shift in chemical composition may significantly affect late season herbivores. 5 These results suggest that aspen chemical resistance mechanisms are ineffective during intense episodic eruptions of outbreak folivores such as the forest tent caterpillar. Variable budbreak phenology may lead to differential susceptibility during less intense outbreak years and, at peak forest tent caterpillar population densities, mechanisms affording tolerance are probably more important than chemical defences.     

Large-scale forest fragmentation increases the duration of tent caterpillar outbreak

I examined historical data (1950–1984) on the duration of outbreaks of the forest tent caterpillar (Malacosoma disstria) in northern Ontario, Canada. Outbreak duration was compared to host tree species dominance and forest structure over large areas of boreal forest partially cleared for agriculture. Abundance of the principal host tree species Populus tremuloides had no consistent effect on duration of outbreak within forest districts, and was negatively correlated with duration of outbreaks among the eight forest districts examined. The amount of forest edge per km² was the best, and most consistent, predictor of the duration of tent caterpillar outbreaks both within individual forest districts and among forest districts. Because forest tent caterpillar populations are driven largely by the impact of parasitoids and pathogens, results here suggest that large-scale increase in forest fragmentation affects the interaction between these natural enemies and forest tent caterpillar. Increased clearing and fragmentation of boreal forests, by agriculture and forestry, may be exacerbating outbreaks of this forest defoliator.     

Nuclear polyhedrosis virus as a biological control agent for Malacosoma americanum (Lepidoptera: Lasiocampidae)

In addition to damaging trees, the eastern tent caterpillar is implicated in early fetal loss and late‐term abortion in horses. In a field study, we evaluated the potential biological control of the caterpillar using eastern tent caterpillar nuclear polyhedrosis virus (ETNPV), a naturally occurring virus that is nearly species‐specific. Egg masses were hatched and second instar larvae were fed virus‐inoculated foliage to propagate the virus in vivo. Then, a viral pesticide was formulated at concentrations of 10⁴, 10⁶ and 10⁸ polyhedral inclusion bodies per ml. The pesticide was applied to foliage on which second, third and fourth instar caterpillars were feeding. When the majority of surviving larvae reached the sixth instar, colonies were collected and the surviving caterpillars counted. Mean numbers of surviving caterpillars per treatment were compared via 95% bootstrap confidence intervals. The data indicate second instar caterpillars were highly susceptible to the virus, but only at the highest concentration tested. Third instar caterpillars were also somewhat susceptible to high virus concentrations, while fourth instar caterpillars were fairly resistant. Our data provide the strongest evidence to date that ETNPV can be propagated, harvested and refined for formulation as a biological control agent for eastern tent caterpillar. Its use on this insect may be merited in circumstances where landowners and managers need to protect trees and horses.     

Adaptation in an insect host–plant pathogen interaction

Selection on parasites to adapt to local host populations may be direct or through other components of the system such as vectors or the food plant on which the parasite is ingested. To test for local adaptation of nucleopolyhedrovirus among island populations of western tent caterpillars, Malacosoma californicum pluviale, we compared virus isolates from three geographically distinct sites with different dominant host plants. Pathogenicity, speed of kill and virus production of each isolate were examined on the three food plants. Virus isolates from the two permanent host populations had the fastest speed of kill on the host plant from which they were isolated. This was not the case for a caterpillar population that goes extinct when populations are regionally low. Virus isolates on some plant species combined rapid speed of kill with high virus yield. Infection of hosts by mixed microparasite populations could facilitate local adaptation in response to differing food plant chemistry.     

Life‐history consequences and disease resistance of western tent caterpillars in response to localised,herbivore‐induced changes in alder leaf quality

1. Plants can respond to herbivore damage with phenotypically plastic changes in quality that negatively affect herbivores and prevent subsequent attack – induced defences. 2. The present study tested whether trees respond to herbivory with localised induction, and whether life‐history traits and disease resistance of an insect herbivore are altered on induced branches of the trees. 3. The influences of localised, within‐branch, herbivore‐induced changes in red alder trees (Alnus rubra Bong.) on fitness characteristics of western tent caterpillars (Malacosoma californicum pluviale Dyar) were evaluated. In the field, randomly selected branches of trees were infested with tent caterpillar larvae and the adjacent branches were maintained as non‐infested controls. In the laboratory, larvae were fed leaves from either induced or non‐induced branches through to adult emergence. A second cohort of larvae was challenged with a viral pathogen to compare their disease susceptibility on induced versus non‐induced foliage. 4. Herbivore‐induced, localised responses of damaged branches reduced leaf quality for growth and the fecundity of female western tent caterpillars, but not that of males. Larvae fed induced leaves had a higher survival overall and a reduced mortality due to unidentified non‐viral pathogens than did their counterparts on non‐induced leaves. However, there was no influence of leaf quality on baculovirus‐induced mortality. 5. These findings suggest that localised induced changes in leaf quality could potentially influence populations of tent caterpillars in contradictory ways by reducing their growth rate and fecundity to a modest degree, while improving their survival and resistance to unidentified non‐viral pathogens to a larger extent.     

Changes in the fecundity of tent caterpillars: a correlated character of disease resistance or sublethal effect of disease?

Over the fluctuation in population density of tent caterpillars, Malacosoma californicum pluviale and M. disstria, fecundity changes from being high at peak density to low for several years during the decline. During the increase phase, fecundity rapidly returns to moderately high levels with a further increase occurring to-ward the end of the increase phase. Two hypotheses which might explain these shifts are that (1) mortality from viral disease which is common during population declines selects for resistant individuals with low fecundity as an associated characteristic, and (2) sublethal viral disease reduces fecundity of moths during population decline. In this study we observed rapid shifts in the frequencies of large and small egg masses and in the mean fecundity between different phases of the population fluctuation. Viral disease was more common in caterpillars from small egg masses of the forest tent caterpillar. These observations are consistent with the hypothesis that sublethal effects of virus reduce the fecundity of moths during the population decline, but high fecundity is quickly restored when disease is rare during the population increase.     

Foliar quality influences tree-herbivore-parasitoid interactions: effects of elevated CO<Subscript>2</Subscript>, O<Subscript>3</Subscript>, and plant genotype

This study examined the effects of carbon dioxide (CO₂)-, ozone (O₃)-, and genotype-mediated changes in quaking aspen (Populus tremuloides) chemistry on performance of the forest tent caterpillar (Malacosoma disstria) and its dipteran parasitoid (Compsilura concinnata) at the Aspen Free-Air CO₂ Enrichment (FACE) site. Parasitized and non-parasitized forest tent caterpillars were reared on two aspen genotypes under elevated levels of CO₂ and O₃, alone and in combination. Foliage was collected for determination of the chemical composition of leaves fed upon by forest tent caterpillars during the period of endoparasitoid larval development. Elevated CO₂ decreased nitrogen levels but had no effect on concentrations of carbon-based compounds. In contrast, elevated O₃ decreased nitrogen and phenolic glycoside levels, but increased concentrations of starch and condensed tannins. Foliar chemistry also differed between aspen genotypes. CO₂, O₃, genotype, and their interactions altered forest tent caterpillar performance, and differentially so between sexes. In general, enriched CO₂ had little effect on forest tent caterpillar performance under ambient O₃, but reduced performance (for insects on one aspen genotype) under elevated O₃. Conversely, elevated O₃ improved forest tent caterpillar performance under ambient, but not elevated, CO₂. Parasitoid larval survivorship decreased under elevated O₃, depending upon levels of CO₂ and aspen genotype. Additionally, larval performance and masses of mature female parasitoids differed between aspen genotypes. These results suggest that host-parasitoid interactions in forest systems may be altered by atmospheric conditions anticipated for the future, and that the degree of change may be influenced by plant genotype.     

Plasticity in Forest Tent Caterpillar Collective Foraging Schedules

Self‐organization can generate synchronized group activity without external triggering cues, and schedules of self‐organized collective activity can vary with environmental conditions. This plasticity can improve group members’ ability to meet their requirements in different environments. In colonial caterpillars, synchronized colony foraging schedules have been postulated to depend either on avoidance of visual predators or on temperature effects on ectotherm physiology. We examine the foraging schedule of forest tent caterpillars (Malacosoma disstria) under different constant conditions to distinguish between these hypotheses. Plasticity in the foraging schedule was tested by keeping colonies under different constant regimes of light and temperature. Digital video and tracking software were used to record the colony's alternation between quiescent and active bouts. The duration and frequency of bouts was compared between treatments. The schedule of synchronized colony activity was not affected by lighting, but it accelerated at higher temperature, because of a decrease in the duration of both active and quiescent bouts. Forest tent caterpillars’ foraging schedule thus depends on the time required to accomplish the tasks of food finding (active bouts) and food processing (quiescent bouts). As caterpillars are ectotherms, locomotion and digestion rates increase at higher temperature and both tasks are accomplished faster. The forest tent caterpillar and the congeneric eastern tent caterpillar (M. americanum) both exhibit self‐organized synchronized collective foraging, but environmental modulation of foraging schedule differs between these species, according to differences in social organization and thermal ecology. Eastern tent caterpillars maintain a fixed foraging schedule under varying temperatures and use the tent to maintain high metabolic rates. In the forest tent caterpillar, flexibility of the foraging schedule in accordance with changes in metabolism lessens the constraints imposed by collective foraging. Synchronous foraging, where entire social groups travel together to and from feeding sites, is thought to have several fitness advantages including improved food finding, recruitment to profitable food sources, anti‐predator defense and group thermoregulation between foraging expeditions.     

Orientation and Dispersal Patterns of the Eastern Tent Caterpillar, <Emphasis Type="Italic">Malacosoma americanum</Emphasis> F. (<Emphasis Type="Italic">Lepidoptera: Lasiocampidae</Emphasis>)

Larvae of the eastern tent caterpillar, Malacosoma americanum, undergo density-dependent dispersal in response to depleted resources. Because these caterpillars have recently been implicated in abortions of pregnant mares (equine Mare Reproductive Loss Syndrome, or MRLS), there is increased interest in managing caterpillar populations, potentially through manipulation of caterpillar dispersal behavior. Consequently, we investigated dispersal patterns of food-deprived eastern tent caterpillars in artificial arenas with respect to distance, direction, and response to visual stimuli. Distance traveled is influenced by time of day, and is strongly correlated with time elapsed. Movement is non-random, and correlates closely with the position of the sun. The pattern is more pronounced with foraging third instars than with penultimate fifth instars. Visual cues appear important in caterpillar orientation, and caterpillars are responsive to vertically oriented, black objects.     

Spillover of tent caterpillar (Malacosoma americanum) herbivory onto willow bioenergy crops in an agricultural landscape

The circumstances and potential for insects to damage perennial bioenergy crops is not well understood in the United States. In this study, we evaluated the spillover and herbivory of eastern tent caterpillars (Malacosoma americanum) from host trees onto short rotation coppice (SRC) willow bioenergy crops (Salix sp.). Host trees were all in the Rosaceae family and included Prunus americana, Prunus virginiana and Malus sp. Willow showed greater leaf herbivory with increasing proximity to a defoliated host tree, suggesting that tent caterpillars spilled‐over to willow after denuding their host. More tent caterpillar herbivory was associated with greater mortality of willow. This study suggests that landscape context and spatial position of host trees is important to the early establishment of a willow bioenergy crop.     

Entomophages et maladies des insectes

Summary Without discussing the problem of the particular role of Epizootics in natural populations, the author examines the different types of relations existing between entomophagous insects and diseases of their hosts. The consequence for parasites and predators of a possible use of pathogenes as a method of Biological Control of pests is examined with more detail in the case ofP. brassicae L. submitted to sprays with a suspension of spores ofB. thuringiensis Berliner. It is shown that two hymenopterous parasites (Apanteles glomeratus L. Anilastus ebeninus Grav). can survive the death of their host, under certain circumstances. The survival ofA. glomeralus is conditionned by its being in the last larval instar on the day of treatment. The consequence of the disease of the host caterpillar is a shortening of the endoparasitic life of the Braconid. A similar effect can be observed whenP. brassicae caterpillars are treated with a contact insecticide, but in this case,Apanteles larvae are killed by residues on their emergence from the host. There is no direct effect of spore suspensions on the larvae of the parasite. It seems thus possible to adopt bacterial control measures preserving a part of the entomophagous populations.

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I.N.R.A., Laboratoire de lutte biologique de La Minière.     

Population dynamics of the beech caterpillar, <Emphasis Type="Italic">Syntypistis punctatella</Emphasis>, and biotic and abiotic factors

The beech caterpillar, Syntypistis punctatella (Motschulsky) (Lepidoptera: Notodontidae), often causes extensive defoliation of beech forests in Japan. Outbreaks have often occurred synchronously among different areas at intervals of 8–11 years. Synchrony of outbreaks was considered to be caused by synchrony of weather. Populations of this insect exhibit periodical dynamics in both outbreak and nonoutbreak areas. Factors that might influence the population dynamics of the beech caterpillar were classified from the point of view of the natural bioregulation com-plex, which includes a coleopteran predator, Calosoma maximowiczi, avian predators, parasitoids, entomopathogenic fungi, and delayed induced defensive response (DIR) of beech trees. Because such periodic population dynamics are believed to be caused by one or more delayed density-dependent factors, delayed density-dependent mortality has been identified as a likely source of population cycles. The DIR and pathogenic diseases showed a high order of density dependence. An infectious pathogen, Cordyceps militaris, was considered to be the most plausible agent responsible for periodic dynamics of the beech caterpillar population because insect diseases were effective in cases in which the S. punctatella population started to decrease without reaching outbreak densities, but DIR was not. Conspicuous defoliation caused by this insect tends to occur at certain elevations, where forests are composed of pure stands of beech trees. I propose three different hypotheses to explain this phenomenon: the diversity–stability hypothesis, the resource concentration hypothesis, and the altitudinal soil nutrient hypothesis. Received: November 20, 1999 / Accepted: August 3, 2000     

Preference and performance of Lepidoptera varies with tree age in juniper woodlands

1. As trees age, they undergo significant physiological and morphological changes. Nevertheless, tree ontogeny and its impacts on herbivores are often overlooked as determinants of plant–herbivore population dynamics and the strength of plant–herbivore interactions. 2. Juniperus (Cupressaceae) is a dominant, long‐lived conifer that serves as the sole host to a specialised assemblage of caterpillars. Over the past 150 years, several juniper species in western North America have expanded their geographic occupancy at local and regional scales, which has resulted in an increase in the number of immature trees on the landscape. Using assays in the laboratory, the effects of tree ontogeny on caterpillar performance and oviposition preference for two juniper specialist caterpillars, Callophrys gryneus (Lycaenidae) and Glena quinquelinearia (Geometridae), were examined. The study considered whether responses to tree ontogeny were consistent across caterpillar species and juniper host species. 3. Tree age was found to be a reliable predictor of caterpillar performance, with caterpillars developing more quickly and growing larger when fed foliage from young trees. Differences in the phytochemical diversity between foliage from trees of different ages might help to explain observed differences in caterpillar performance. Interestingly, the specialist butterfly, C. gryneus, displayed an oviposition preference for foliage from old‐growth Juniperus osteosperma trees, despite the fact that larvae of this species performed poorly on older trees. 4. It is concluded that young juniper trees are an important resource for the specialised Lepidopteran community and that tree ontogeny is an important component of intraspecific variation, which contributes to the structure of plant–herbivore communities.     

Previous herbivore attack of red alder may improve food quality for fall webworm larvae

Summary Three hypotheses of insect-plant interactions were tested by rearing fall webworm larvae in the laboratory on foliage from red alder trees with different histories of western tent caterpillar herbivory. Fall webworm larvae raised on foliage from trees which had been attacked previously for two summers by moderate densities of western tent caterpillars grew faster and attained heavier pupal weights than did those fed foliage from unattacked trees. This contradicts the hypothesis that moderate levels of previous herbivory induces the production of plant defensive chemicals in red alders. Growth of webworms, when fed foliage from unattacked trees adjacent to alders that were attacked by fall webworm larvae, was the same as when fed foliage from trees isolated by distance from attacked trees. This contradicts the hypothesis that attacked trees stimulate the production of defensive chemicals in neigh-boring trees. Young and mature alder foliage was equally good for fall webworm growth and survival, and foliage from trees heavily attacked by both fall webworm and western tent caterpillars for three years produced slow growth rates and small pupal sizes. This supports the hypothesis that continued heavy insect attack can cause the deterioration of the food quality of attacked trees.