Western Tent Caterpillar Research Articles

Warmer temperatures reduce the transmission of a virus in a gregarious forest insect.

Understanding how climate warming will influence species interactions is a key question in ecology and predicting changes in the prevalence of disease outbreaks is particularly challenging. Ectotherms are likely to be more influenced by climatic changes as temperature governs their growth, feeding, development, and behavior. We test the hypothesis that pathogen transmission and host mortality will increase at warmer temperatures using a cyclic forest insect, the western tent caterpillar (WTC), Malacosoma californicum pluviale, and its baculovirus. The virus causes population declines at peak host density. WTC are gregarious and clustering is predicted to increase the risk of within family infection; however, how temperature influences this has not been examined. We investigated the impact of temperature on different components of the transmission process in order to pinpoint the possible mechanisms involved. In the laboratory, leaf consumption increased linearly with rising temperature between 15 and 30°C. Insects died more rapidly from virus infection as temperature increased, but this did not translate into differences in the production of viral transmission stages. To examine the influence of temperature on virus transmission, we created a temperature difference between two greenhouses containing potted red alder trees, Alnus rubra. The cooler greenhouse (mean 19.5°C) was roughly similar to ambient temperatures in the field, while the warmer greenhouse was 10°C higher (mean 29°C). As predicted, both larval movement and feeding were higher at the warmer temperature, while the likelihood of the preinfected, inoculum larvae dying on the tents was twice as high in the cooler greenhouse. This resulted in increased virus mortality and a higher transmission parameter under cooler conditions. Therefore, we suggest that, contrary to our prediction, the reduced movement of infected larvae at colder temperatures increased the risk of infection in these gregarious insects and had a greater impact on virus transmission than the increased activity of the susceptible larvae in warmer conditions. Long-term population data from the field, however, show no relationship between temperature and infection levels, suggesting that local changes in virus transmission might not scale up to population infection levels.

Spatial Habitat Structure Assembles Willow-Dependent Communities across the Primary Successional Watersheds of Mount St. Helens, USA

The eruption of Mount St. Helens in 1980 resulted in a cataclysmic restructuring of its surrounding landscapes. The Pumice Plain is one of these landscapes, where tree species such as Sitka willow (Salix sitchensis) and their dependent communities have been established along newly-formed streams. Thus, the study of these dependent communities provides a unique and rare opportunity to investigate factors influencing metacommunity assembly during true primary succession. We analyzed the influence of landscape connectivity on metacommunity assembly through a novel application of circuit theory, alongside the effects of other factors such as stream locations, willow leaf chemistry, and leaf area. We found that landscape connectivity structures community composition on willows across the Pumice Plain, where the least connected willows favored active flyers such as the western tent caterpillar (Malacosoma fragilis) or the Pacific willow leaf beetle (Pyrrhalta decora carbo). We also found that multiple levels of spatial habitat structure linked via landscape connectivity can predict the presence of organisms lacking high rates of dispersal, such as the invasive stem-boring poplar weevil (Cryptorhynchus lapathi). This is critical for management as we show that the maintenance of a heterogeneous mixture of landscape connectivity and resource locations can facilitate metacommunity dynamics to promote ecosystem function and mitigate the influences of invasive species.

Bottom-up and top-down pressures mediate competition between two generalist insects.

The effects of competition can have far-reaching consequences for individuals, populations, and communities and therefore we should strive toward a deeper understanding of competitive interactions. In some cases, dietary generalists may be predicted to experience weak competition effects because of their ability to use a wide range of host plants. However, competition between insects frequently occurs indirectly, which can hinder insects' abilities to avoid competitive interactions. Therefore, competition may be as strong among dietary generalists as among dietary specialists. Yet competition between insects that are dietary generalists is infrequently studied. We tested for evidence of competitive interactions between two common, temporally separated, generalist insects: the western tent caterpillar (Malacosoma californicum), which feeds early in the season, and the fall webworm (Hyphantria cunea), which feeds later in the season. Both species frequently use a common host plant species (chokecherry) as a preferred host at our field sites. We tested the relative strength of bottom-up effects resulting from competitive interactions between these two generalists with laboratory-rearing trials at the relevant time of year for each insect. We recorded three common fitness measures (development time, pupal mass, and survival) for caterpillars reared on chokecherry with no damage from either of our focal species, with tent caterpillar damage, and with fall webworm damage. To test the strength of top-down pressures on fall webworm larval fitness and any potential interactions with bottom-up effects, we reared larvae in the field either exposed to or protected from predators on host plants that either did or did not have tent caterpillars feeding on them earlier in the season. We found evidence of bottom-up fitness effects on tent caterpillars and top-down and bottom-up fitness effects on fall webworms confirming that tent caterpillars and fall webworms compete indirectly. Tent caterpillars had lower pupal mass when reared on leaves from shrubs damaged by fall webworms. Fall webworms had lower pupal mass and longer development time when reared on leaves from shrubs damaged by tent caterpillars. In field trials, fall webworms reared on shrubs damaged by tent caterpillars had a lower survival and pupal mass. We show evidence of indirect competition in temporally separated generalists through leaf quality (bottom-up effects) and natural enemies (top-down effects).

Factors Affecting Foliar Oregonin and Condensed Tannin in Red Alder (Alnus rubra Bong.): Phytochemicals Implicated In Defense Against Western Tent Caterpillar (Malacosoma californicum Packard).

Past work shows a significant negative correlation between foliar oregonin concentration and western tent caterpillar (Malacosoma californicum Packard) feeding on red alder (Alnus rubra Bong.). Above an oregonin threshold of 20% leaf dry weight, little feeding by caterpillars is observed. Concentrations of defensive chemicals are influenced by plant genotype, environmental conditions, insect feeding, and the interactions of these factors. Our objective was to measure the effects of nitrogen (N) availability and wounding on foliar oregonin and condensed tannin concentrations in red alder genotypes. One-year-old seedlings from 100 half-sib red alder families were treated with two levels of ammonium nitrate (NH4NO3) for two growing seasons in a common garden. In the second year, leaves from 50 families from the fertilization experiment were used in a bioassay feeding experiment to determine the effects of N fertilization and genotype on WTC damage, and to identify a subset of 20 families with a range of damage to analyze for phytochemical composition. In separate experiments, wound-induction treatments were conducted outdoors and, in a greenhouse using the N treated trees in their third and fourth year, respectively. Foliar condensed tannin, oregonin and N concentrations were measured and ranked among the plant genotypes, and between the two N treatments and two wounding treatments. Results showed that oregonin and condensed tannin concentrations varied among the alder genotypes. Leaf N concentration was negatively correlated with concentration of oregonin. Neither of the measured phenolic compounds responded to wounding. The results suggest that red alder foliar oregonin and condensed tannin are likely constitutive defenses that are largely determined by genotype, and that the negative correlation of defense compounds with plant internal N status holds in this N-fixing tree.

Anti-Herbivore Activity of Oregonin, a Diarylheptanoid Found in Leaves and Bark of Red Alder (Alnus rubra).

Plants synthesize a wide range of bioactive secondary metabolites to defend against pests and pathogens. Red alder (Alnus rubra) bark, root, and leaf extract have a long history of use in traditional medicine and hygiene. Diarylheptanoids, especially oregonin ((5S)-1,7-bis(3,4-dihydroxyphenyl)-5-(β-D-xylopyranosyloxy)-heptan-3-one), have been identified as major bioactive constituents. Diarylheptanoids have become a focus of research following reports of their antioxidant, antifungal, and anti-cancer activities. Recent data suggest that high oregonin concentration is associated with resistance of red alder leaves to western tent caterpillar (Malacosoma californicum) defoliation. Here we test effects of this compound directly on leaf-eating insects. Purified oregonin was examined in insect choice and toxicity tests using lepidopteran caterpillars. The compound exhibited significant anti-feedant activity against cabbage looper (Trichoplusia ni), white-marked tussock moth (Orgyia leucostigma), fall webworm (Hyphantria cunea), and M. californicum at concentrations corresponding to oregonin content of the most resistant alder clones in previous experiments. Toxicity tests were carried out with cabbage looper larvae only, but no contact or ingested toxicity was detected. Our results suggest that oregonin at levels found in red alder leaves early in the growing season may contribute to protecting red alder from leaf-eating insects.

Red alder defense mechanisms against western tent caterpillar defoliation

Red alder (Alnus rubra Bong.) is a tree of high economic and ecological importance, yet is subject to severe defoliation during episodic outbreaks of tent caterpillars (Malacosoma spp.). We evaluated variation in western tent caterpillar (Malacosoma californicum Packard, 1864) (WTC) resistance among and within red alder populations and clones and investigated potential defense mechanisms. Bioassay feeding trials were conducted with WTC on 20 red alder clones from 10 provenances (two clones per provenance). Phenology and quality of red alder leaves were analyzed to determine whether bud burst, leaf chemical content, water content, or physical traits are determinants of WTC preference. Alder clones differed in percent leaf area eaten by WTC and in leaf defense traits. The concentrations of total phenolics, condensed tannin, and the diarylheptanoid oregonin were negatively correlated with the percent leaf area eaten by caterpillars. In addition, a potential threshold was observed above which the concentration of each of the chemicals appeared to reduce WTC feeding. In particular, foliar concentrations of oregonin >20% of leaf dry mass were consistently associated with reduced feeding. The effects of oregonin concentration in red alder leaves on tent caterpillar feeding is a novel finding.

The Role of Enemy-Mediated Competition in Determining Fitness of a Generalist Herbivore

The influence of indirect competition on insect communities is not well understood. We studied how the fitness of a gregarious lepidopteran species, fall webworm, Hyphantria cunea (Drury), is affected by another gregarious caterpillar species, western tent caterpillar, Malacosoma californicum (Packard). Fall webworm and tent caterpillars both feed on chokecherry, Prunus virginiana L., and create silken structures in which they reside until just prior to pupation. Preliminary observations suggested that predators and parasitoids reside in tent caterpillar tents after tents have been abandoned by the caterpillars, thus increasing the presence of enemies near tents. We hypothesized that when reared near tent caterpillar tents, fall webworms are less fit because of increased predation from nearby enemies (enemy-mediated competition). We tested our hypothesis at two field sites by placing fall webworm larvae in the field with or without a tent caterpillar tent nearby. We compared differences in survival rates, parasitism rates, and fitness between the larvae reared with and without tents nearby. At one field site, webworm larvae near tent caterpillar tents disappeared ∼30% faster than did larvae without tents nearby. Although survival or fitness did not differ significantly between larvae reared with or without tents nearby, distance separating fall webworm larvae from a tent caterpillar tent might be important. Our results suggested that enemy-mediated competition might affect the fitness of fall webworm larvae and should be investigated further.

Time-lagged intraspecific competition in temporally separated cohorts of a generalist insect.

Competition can have far-reaching consequences for insect fitness and dispersion. Time-lagged interspecific competition is known to negatively affect fitness, yet time-lagged intraspecific competition is rarely studied outside of outbreak conditions. We tested the impact of competition between larval cohorts of the western tent caterpillar (Malacosoma californicum) feeding on chokecherry (Prunus virginiana). We reared larvae on host plants that either had or did not have feeding damage from tent caterpillars the previous season to test the bottom-up fitness effects of intraspecific competition. We measured host-plant quality to test potential mechanisms for bottom-up effects and conducted field oviposition surveys to determine if female adult tent caterpillars avoided host plants with evidence of prior tent caterpillar presence. We found that time-lagged intraspecific competition impacted tent caterpillar fitness by reducing female pupal mass, which is a predictor of lifetime fitness. We found that plants that had been fed upon by tent caterpillars the previous season had leaves that were significantly tougher than plants that had not been fed upon by tent caterpillars, which may explain why female tent caterpillars suffered reduced fitness on these plants. Finally, we found that there were fewer tent caterpillar egg masses on plants that had tent caterpillars earlier in the season than plants without tent caterpillars, which suggests that adult females avoid these plants for oviposition. Our results confirm that intraspecific competition occurs among tent caterpillars and suggests that time-lagged intraspecific competition has been overlooked as an important component of insect fitness.

Phylloplane bacteria increase the negative impact of food limitation on insect fitness

1. When populations of herbivorous insects increase in density, they can alter the quantity or quality of their food. The impacts of diet‐related stressors on insect fitness have been investigated singly, but not simultaneously.2. Foliage quantity and quality of red alder, Alnus rubra, were manipulated together with the presence of non‐entomopathogenic phylloplane bacteria to investigate their impacts, singly and in combination, on survival, pupal mass, growth rate, fecundity and egg quality of a cyclic forest insect, the western tent caterpillar, Malacosoma californicum pluviale.3. Food limitation (half food) had strong negative impacts on all life‐history traits. When the larvae were fed continuously, however, neither ingesting phylloplane bacteria nor eating leaves from damaged branches (induced foliage) affected survival. In the half‐food treatment, ingesting bacteria further increased mortality, while feeding on induced foliage improved survival.4. Growth rate and pupal mass of both sexes were reduced for larvae with food limitation compared with continuously fed insects and this was exacerbated when the larvae also ate bacteria‐treated leaves. A combination of bacteria and induced foliage also reduced larval growth rate by 5% in the full‐food treatment.5. Fecundity (eggs per egg mass) was 2.7 times greater in full‐food than in food‐limited treatments but neither phylloplane bacteria nor plant induction had an effect. Insects fed induced foliage produced smaller eggs. Overall, there was no evidence of a three‐way interaction between the three stressors, although there were negative synergistic effects, primarily between food limitation and the ingestion of phylloplane bacteria.

Performance of Western Tent Caterpillar (Malacosoma californicum) on Two Common Host Plants, Including a New Host Plant Record

ABSTRACT. Tent caterpillars are generalists across their full host range, but display local host plant preferences. We present evidence for a new host plant record, wax currant (Ribes cereum), for western tent caterpillars (Malacosoma californicum) along the Colorado Front Range. We tested the suitability of wax currant as a host plant for western tent caterpillars as compared to chokecherry (Prunus virginiana), an abundant and commonly used host plant. We measured the density of tent caterpillar tents in areas where both host plants occur to assess host plant use. We reared tent caterpillar larvae on both host plants and measured fitness effects due to host plant quality (survival, pupal mass) and natural enemies (parasitism). We did not find a relationship between host plant abundance and use by tent caterpillars and found no evidence for a preference for either host plant. We found that western tent caterpillars do not differ in pupal mass when reared on chokecherry and on wax currant in a laboratory s...

Ecology and evolution of pathogens in natural populations of Lepidoptera.

Pathogens are ubiquitous in insect populations and yet few studies examine their dynamics and impacts on host populations. We discuss four lepidopteran systems and explore their contributions to disease ecology and evolution. More specifically, we elucidate the role of pathogens in insect population dynamics. For three species, western tent caterpillars, African armyworm and introduced populations of gypsy moth, infection by nucleopolyhedrovirus (NPV) clearly regulates host populations or reduces their outbreaks. Transmission of NPV is largely horizontal although low levels of vertical transmission occur, and high levels of covert infection in some cases suggest that the virus can persist in a nonsymptomatic form. The prevalence of a mostly vertically transmitted protozoan parasite, Ophryocystis elektroscirrha, in monarch butterflies is intimately related to their migratory behaviour that culls highly infected individuals. Virulence and transmission are positively related among genotypes of this parasite. These systems clearly demonstrate that the interactions between insects and pathogens are highly context dependent. Not only is the outcome a consequence of changes in density and genetic diversity: environmental factors, particularly diet, can have strong impacts on virulence, transmission and host resistance or tolerance. What maintains the high level of host and pathogen diversity in these systems, however, remains a question.

The effects of experimental warming on the timing of a plant-insect herbivore interaction.

The phenology of many species is shifting in response to climatic changes, and these shifts are occurring at varying rates across species. This can potentially affect species' interactions and individual fitness. However, few studies have experimentally tested the influence of warming on the timing of species interactions. This is an important gap in the literature given the potential for different direct and indirect effects of temperature via phenological change. Our aim was to test the effects of warming on the western tent caterpillar (Malacosoma californicum pluviale). In addition to the direct effects of warming, we considered the two primary indirect effects mediated by warming-driven changes in its host plant, red alder (Alnus rubra): changes in resource availability due to phenological mismatch (i.e. changes in the relative timing of the interaction), and changes in resource quality associated with leaf maturation. We experimentally warmed egg masses and larvae of the western tent caterpillar placed on branches of red alder in the field. Warming advanced the timing of larval but not leaf emergence. This led to varying degrees of phenological mismatch, with larvae emerging as much as 25 days before to 10 days after the emergence of leaves. Even the earliest-emerging larvae, however, had high survival in the absence of leaves for up to 3 weeks, and they were surprisingly resistant to starvation. In addition, although warming created phenological mismatch that initially slowed the development of larvae that emerged before leaf emergence, it accelerated larval development once leaves were available. Therefore, warming had no net effect on our measures of insect performance. Our results demonstrate that the indirect effects of warming, in creating phenological mismatch, are as important to consider as the direct effects on insect performance. Although future climatic warming might influence plants and insects in different ways, some insects may be well adapted to variation in the timing of their interactions.

Genetic similarity of island populations of tent caterpillars during successive outbreaks.

Cyclic or fluctuating populations experience regular periods of low population density. Genetic bottlenecks during these periods could give rise to temporal or spatial genetic differentiation of populations. High levels of movement among increasing populations, however, could ameliorate any differences and could also synchronize the dynamics of geographically separated populations. We use microsatellite markers to investigate the genetic differentiation of four island and one mainland population of western tent caterpillars, Malacosoma californicum pluviale, in two periods of peak or pre-peak density separated by 8 years. Populations showed high levels of genetic variation and little genetic differentiation either temporally between peaks or spatially among sites. Mitochondrial haplotypes were also shared between one island population and one mainland population in the two years studied. An isolation-by-distance analysis showed the FST values of the two geographically closest populations to have the highest level of differentiation in both years. We conclude that high levels of dispersal among populations maintain both synchrony of population dynamics and override potential genetic differentiation that might occur during population troughs. As far we are aware, this is the first time that genetic similarity between temporally separated population outbreaks in insects has been investigated. A review of genetic data for both vertebrate and invertebrate species of cyclic animals shows that a lack of spatial genetic differentiation is typical, and may result from high levels of dispersal associated with fluctuating dynamics.

Tent caterpillars are robust to variation in leaf phenology and quality in two thermal environments

The synchrony between emergence of spring-active, insect herbivores and the budburst of their host plants could be affected by warming temperatures with influences on the availability and quality of foliage as it undergoes physical and chemical changes. This can affect the growth and survival of insects. Here, we used sun-exposed and shaded trees to determine whether the synchrony between egg hatch of western tent caterpillar, Malacosoma californicum pluviale Dyar (Lepidoptera:Lasiocampidae) and budburst of its host red alder, Alnus rubra Bongard (Betulaceae)changes with different thermal environments (temperature and light together). To explore the potential outcome of a shift in phenological synchrony, we used laboratory assays of larval growth and survival to determine the effect of variation in young, youthful and mature leaves from sun-exposed and shaded trees. While the average higher temperature of sun-exposed trees advanced the timing of budburst and egg hatch, synchrony was not disrupted. Leaf quality had no significant influence on growth or survival in the laboratory for early instars reared as family groups. Later instar larvae, however, performed best on mature leaves from sun-exposed trees. The robust relationship between leaf and larval development of western tent caterpillars suggests that warming climates may not have a strong negative impact on their success through shifts in phenological synchrony, but might influence other aspects of leaf quality and larval condition.

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.

Multiple mating and family structure of the western tent caterpillar, Malacosoma californicum pluviale: impact on disease resistance.

BackgroundLevels of genetic diversity can strongly influence the dynamics and evolutionary changes of natural populations. Survival and disease resistance have been linked to levels of genetic diversity in eusocial insects, yet these relationships remain untested in gregarious insects where disease transmission can be high and selection for resistance is likely to be strong.Methodology/Principal FindingsHere we use 8 microsatellite loci to examine genetic variation in 12 families of western tent caterpillars, Malacosoma californicum pluviale from four different island populations to determine the relationship of genetic variability to survival and disease resistance. In addition these genetic markers were used to elucidate the population structure of western tent caterpillars. Multiple paternity was revealed by microsatellite markers, with the number of sires estimated to range from one to three per family (mean ± SE = 1.92±0.23). Observed heterozygosity (HO) of families was not associated to the resistance of families to a nucleopolyhedrovirus (NPV) (r = 0.161, F1,12 = 0.271, P = 0.614), a major cause of mortality in high-density populations, but was positively associated with larval survival (r = 0.635, F1,10 = 5.412, P = 0.048). Genetic differentiation among the families was high (FST = 0.269, P<0.0001), and families from the same island were as differentiated as were families from other islands.Conclusion/SignificanceWe have been able to describe and characterize 8 microsatellite loci, which demonstrate patterns of variation within and between families of western tent caterpillars. We have discovered an association between larval survival and family-level heterozygosity that may be relevant to the population dynamics of this cyclic forest lepidopteran, and this will be the topic of future work.

The effect of food limitation on immunity factors and disease resistance in the western tent caterpillar

Epizootics of nucleopolyhedrovirus characterize declines of cyclic populations of western tent caterpillars, Malacosoma pluviale californicum. In field populations, infection can be apparently lacking in one generation and high in the next. This may suggest an increase in the susceptibility to infection of larvae at peak density or the triggering of a vertically transmitted virus. Here, we test the hypothesis that reduced food availability, as may occur during population outbreaks of tent caterpillars, influences the immunocompetence of larvae and increases their susceptibility to viral infection. We compared immunity factors, hemolymph phenoloxidase and hemocyte numbers, and the susceptibility to nucleopolyhedroviral infection of fifth instar larvae that were fully or partially fed as fourth instars. To determine if maternal or transgenerational influences occurred, we also determined the susceptibility of the offspring of the treated parents to viral infection. Food limitation significantly reduced larval survival, development rate, larval and pupal mass, moth fecundity and levels of hemolymph phenoloxidase, but not the numbers of hemocytes. Neither the food-reduced larvae nor their offspring were more susceptible to viral infection and were possibly even less susceptible at intermediate viral doses. Food reduction did not activate latent or covert viral infection of larvae as might be expected as a response to stress. We conclude that reducing the food intake of fourth instar larvae to an extent that had measurable and realistic impacts on their life history characteristics was not translated into increased susceptibility to viral infection.

Within and between population variation in disease resistance in cyclic populations of western tent caterpillars: a test of the disease defence hypothesis

1. Epizootics of nucleopolyhedrovirus (NPV) are an obvious component of the population fluctuations of several species of temperate forest Lepidoptera, including the western tent caterpillar, Malacosoma californicum pluviale (Dyer). An observed relationship between epizootics and the subsequent reduction in fecundity of populations led to the formulation of the disease defence hypothesis. This hypothesis predicts that viral epizootics in peak populations select for more resistant moths and that their reduced fecundity in declining populations reflects a cost of disease resistance. 2. To test the disease defence hypothesis, we carried out bioassays to measure the variation in larval resistance to NPV infection for families of western tent caterpillars from four spatially distinct populations over 3 years of peak and declining host densities. 3. Each female moth lays a single egg mass and larvae are gregarious and remain together through development. We found that the resistance to disease of larvae within families was not related to the number of eggs in the mass from which they hatched (the fecundity of their mother). 4. Disease resistance of larvae varied among populations and over time in a manner consistent with selection for resistance. One population that had not experienced a strong viral epizootic during the last population decline was more susceptible to infection in the first year of the study. Larvae from a second population that experienced an early epizootic became significantly more resistant. The resistance of two other populations increased slightly before the viral epizootic occurred in the field however, and thus could not be explained by selection. 5. As population densities declined from peak density, the background mortality of larvae increased and the fecundity of moths decreased. This indicates a general deterioration in the quality of field populations of tent caterpillars associated with the declining populations. 6. Although some evidence suggests that viral epizootics can select for increased resistance of field populations of tent caterpillars, the general deterioration in quality, elevated background mortality, and the reduced fecundity after the epizootic are stronger influences on the population decline. These are possibly related to sublethal viral infection.

Adaptation in an insect host–plant pathogen interaction

AbstractSelection 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.

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.