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.     

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.     

Parasitoids on the loose – experimental lack of support of the parasitoid movement hypothesis

Forest fragmentation can disrupt important ecological processes both within and among species. Of particular interest is the extent to which fragmentation decouples economically important pest species from their parasites. In an influential paper, Roland and Taylor (1997) advanced the hypothesis that forest fragmentation at a relatively fine spatial scale would disrupt the ability of parasitoids to aggregate in response to host density, thereby facilitating the build-up of host populations at isolated sites. Our study provides the first experimental test of this notion. We augmented existing densities of forest tent caterpillar individuals at two types of sites: isolated forest fragments and continuous tracts of forest. Of the four parasitoid species studied by Roland and Taylor (1997), two were encountered in our experiment. For the sarcophagid fly Arachnidomyia aldrichi and the tachinid Carcelia malacosomae, we found no difference in parasitism rates among forest fragments and continuous forest tracts. A third tachinid species unique to this study, Lespesia frenchii , is known to be a broad generalist with respect to host choice. In this species, we observed significantly elevated parasitism in forest fragments, and higher parasitism at the forest edge than within the forest. All of these findings suggest complex effects of forest fragmentation on parasitoid movement, and warn against broad generalizations from observational studies. Forest fragmentation may affect movement patterns in a multitude of ways, significantly obscuring or modifying the simplest mechanisms proposed by Roland and Taylor (1997).     

Macrogeographic clines in fecundity, reproductive allocation, and offspring size of the forest tent caterpillar Malacosoma disstria

1. The fecundity of the forest tent caterpillar varies considerably across its geographic range. Field data indicate that populations in the southern United States (Gulf States) produce nearly twice as many eggs as females from Canada or the Lake States, with little or no difference in the size of adult females. 2. In controlled rearing experiments, female forest tent caterpillar from the southern United States (Louisiana) had much larger clutch sizes than same sized females from northern populations in Michigan or Manitoba, Canada. Increased fecundity in Louisiana females was achieved through a significant reduction in egg size and a concomitant increase in the allocation of resources to egg production. 3. Comparison of 10 forest tent caterpillar populations spanning a 27° latitudinal gradient, validated the results of detailed comparisons among the three populations above by confirming the strong negative correlation between latitude and clutch size. 4. Neonate forest tent caterpillars from Manitoba were significantly larger than larvae from either Michigan or Louisiana. Michigan larvae were intermediate in size. It is postulated that large neonates are advantageous in thermally limiting environments. More than three times as many degree‐days are available to Louisiana neonates during the first 2 weeks after hatching. A consistently favourable climate during the vulnerable post‐hatching period may have allowed the evolution of larger clutches at the expense of neonate size in southern populations.     

How parasitism affects critical patch-size in a host-parasitoid model: application to the forest tent caterpillar

Habitat structure has broad impacts on many biological systems. In particular, habitat fragmentation can increase the probability of species extinction and on the other hand it can lead to population outbreaks in response to a decline in natural enemies. An extreme consequence of fragmentation is the isolation of small regions of suitable habitat surrounded by a large region of hostile matrix. This scenario can be interpreted as a critical patch-size problem, well studied in a continuous time framework, but relatively new to discrete time models. In this paper we present an integrodifference host-parasitoid model, discrete in time and continuous in space, to study how the critical habitat-size necessary for parasitoid survival changes in response to parasitoid life history traits, such as emergence time. We show that early emerging parasitoids may be able to persist in smaller habitats than late emerging species. The model predicts that these early emerging parasitoids lead to more severe host outbreaks. We hypothesise that promoting efficient late emerging parasitoids may be key in reducing outbreak severity, an approach requiring large continuous regions of suitable habitat. We parameterise the model for the host species of the forest tent caterpillar Malacosoma disstria Hbn., a pest insect for which fragmented landscape increases the severity of outbreaks. This host is known to have several parasitoids, due to paucity of data and as a first step in the modelling we consider a single generic parasitoid. The model findings are related to observations of the forest tent caterpillar offering insight into this host-parasitoid response to habitat structure.     

Gall wasps and their parasitoids in cork oak fragmented forests

Abstract.  1. This paper explores the potential effects of host-plant fragmentation on cork oak gall wasp populations (Cynipidae, Hymenoptera) and on their predators, lethal inquilines, and parasitoids. To address this objective, galls were collected across a gradient of cork oak ( Quercus suber ) forest fragmentation in the East Pyrenees (Albera, Spain), and they were incubated to obtain the parasitism rates.
2. Two hypotheses were tested: (1) Host-plant fragmentation may induce a decline in gall wasp populations because of area and isolation effects on local extinction and dispersal; as a consequence of that, parasitoids may decline even more strongly in fragmented habitats than their prey. (2) Host-plant fragmentation may cause a decline in gall wasp parasitoid populations that, in turn, can lead to an ecological release in their prey populations.
3. Among the eight cork oak gall wasps sampled in the study area of Albera, the gall abundances of three species ( Callirhytis glandium , Callirhytis rufescens , and Andricus hispanicus ) were significantly related to forest fragmentation. The overall abundance of gall wasps was affected by a radius of ≈ 890 m surrounding landscape, presenting constant abundances with forest loss until forest cover is reduced at ≈ 40%; below that value the abundance increased rapidly. Three inquilines and 23 parasitoids species were recorded after gall incubation. In 25 cases, species of inquilines and parasitoids were newly recorded for the corresponding host in the Iberian peninsula.
4. Although the overall parasitism rate was high (1.1), it was uncorrelated with fragmentation and with overall cynipid abundance. These results indicate that host-plant fragmentation was correlated with higher abundance of gall wasps, whereas the parasitism rate could not explain this hyper-abundance in small forest fragments.     

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.     

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.     

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.     

The impact of parasitoid emergence time on host–parasitoid population dynamics

We investigate the effect of parasitoid phenology on host–parasitoid population cycles. Recent experimental research has shown that parasitized hosts can continue to interact with their unparasitized counterparts through competition. Parasitoid phenology, in particular the timing of emergence from the host, determines the duration of this competition. We construct a discrete-time host–parasitoid model in which within-generation dynamics associated with parasitoid timing is explicitly incorporated. We found that late-emerging parasitoids induce less severe, but more frequent, host outbreaks, independent of the choice of competition model. The competition experienced by the parasitized host reduces the parasitoids’ numerical response to changes in host numbers, preventing the ‘boom-bust’ dynamics associated with more efficient parasitoids. We tested our findings against experimental data for the forest tent caterpillar (Malacosoma disstria Hübner) system, where a large number of consecutive years at a high host density is synonymous with severe forest damage.     

Can caterpillar density or host-plant quality explain host-plant-related parasitism of a generalist forest caterpillar assemblage?

Herbivore–carnivore interactions are influenced by the plants on which herbivores feed. Accordingly, dietary generalist herbivores have been shown to experience differential risk of mortality from carnivores on different host-plant species. Here, we investigate whether caterpillar density and host-plant quality play a role in driving variation in generalist forest caterpillar mortality from insect parasitoids using a large-scale, multi-year observational study. A total of 4,500 polyphagous caterpillars were collected from eight host-tree species in Connecticut deciduous forests over 5 years, and frequencies of mortality from insect parasitoids (flies and wasps) were compared across the eight host-plant species for the entire generalist caterpillar assemblage (76 species). Separate comparisons were made using seven numerically dominant generalist species, allowing us to account for variation in caterpillar species-specific parasitism risk. We find significant variation in parasitism frequencies of generalist caterpillars across the eight host-plant species when accounting for variation in caterpillar density. We find no support for an influence of caterpillar density on parasitism and no clear evidence for an effect of host-plant quality on parasitism. Therefore, the results of this study discount the hypotheses that variation in caterpillar density and host-plant quality are responsible for variation in parasitism frequencies across host-plant species. Instead, our findings point to other plant-related characteristics, such as plant-derived parasitoid attractants, which may have robust, community-wide effects.     

Leaf age effects of elevated CO2-grown white oak leaves on spring-feeding lepidopterans

Folivorous insect responses to elevated CO₂-grown tree species may be complicated by phytochemical changes as leaves age. For example, young expanding leaves in tree species may be less affected by enriched CO₂-alterations in leaf phytochemistry than older mature leaves due to shorter exposure times to elevated CO₂ atmospheres. This, in turn, could result in different effects on early vs. late instar larvae of herbivorous insects. To address this, seedlings of white oak (Quercus alba L.), grown in open-top chambers under ambient and elevated CO₂, were fed to two important early spring feeding herbivores; gypsy moth (Lymantria dispar L.), and forest tent caterpillar (Malacosoma disstria Hübner). Young, expanding leaves were presented to early instar larvae, and older fully expanded or mature leaves to late instar larvae. Young leaves had significantly lower leaf nitrogen content and significantly higher total nonstructural carbohydrate:nitrogen ratio as plant CO₂ concentration rose, while nonstructural carbohydrates and total carbon-based phenolics were unaffected by plant CO₂ treatment. These phytochemical changes contributed to a significant reduction in the growth rate of early instar gypsy moth larvae, while growth rates of forest tent caterpillar were unaffected. The differences in insect responses were attributed to an increase in the nitrogen utilization efficiency (NUE) of early instar forest tent caterpillar larvae feeding on elevated CO₂-grown leaves, while early instar gypsy moth larval NUE remained unchanged among the treatments. Later instar larvae of both insect species experienced larger reductions in foliage quality on elevated CO₂-grown leaves than earlier instars, as the carbohydrate:nitrogen ratio of leaves substantially increased. Despite this, neither insect species exhibited changes in growth or consumption rates between CO₂ treatments in the later instar. An increase in NUE was apparently responsible for offsetting reduced foliar nitrogen for the late instar larvae of both species.     

Spatial and temporal variation in the parasitoid assemblage of an exophytic polyphagous caterpillar

Abstract 1. Over 3400 larvae of the polyphagous ground dwelling arctiid Grammia geneura were sampled and reared over seven generations in order to characterise its parasitoid assemblage and examine how and why this assemblage varies over time and space at a variety of scales.
2. The total parasitoid assemblage of 14 species was dominated both in diversity and frequency by relatively polyphagous tachinid flies.
3. Both the composition of the parasitoid assemblage and frequency of parasitism varied strikingly among and within sampling sites, seasons, and years.
4. Overall rates of parasitism increased consistently over the duration of caterpillar development.
5. Within sampling sites, parasitism rates were non-random with respect to habitat structure and caterpillar behaviour for the most abundant parasitoid species.
6. The large variability in parasitoid assemblage structure over space and time in this system may be a function of local host population abundance, habitat-specific parasitism, and indirect interactions between G. geneura and other Macrolepidoptera through shared oligophagous and polyphagous parasitoids.     

Delayed density-dependent parasitism of eggs and pupae as a contributor to the cyclic population dynamics of the autumnal moth

Many populations of forest Lepidoptera exhibit 10-year cycles in densities, with impressive outbreaks across large regions. Delayed density-dependent interactions with natural enemies are recognized as key factors driving these cyclic population dynamics, but emphasis has typically been on the larval stages. Eggs, pupae and adults also suffer mortality from predators, parasitoids and pathogens, but little is known about possible density relationships between mortality factors and these non-feeding life stages. In a long-term field study, we experimentally deployed autumnal moth (Epirrita autumnata) eggs and pupae to their natural enemies yearly throughout the 10-year population cycle in northern Norway. The abundance of another geometrid, the winter moth (Operophtera brumata), increased in the study area, permitting comparisons between the two moth species in predation and parasitism. Survival of autumnal moth eggs and pupae was related to the moth abundance in an inverse and delayed manner. Egg and pupal parasitoids dominated as density-dependent mortality factors and predicted the subsequent growth rate of the host population size. In contrast, effects of egg and pupal predators were weakly density dependent, and generally predation remained low. Parasitism rates did not differ between the autumnal and winter moth pupae, whereas predators preferred winter moth pupae over those of the autumnal moth. We conclude that parasitism of the autumnal moth by egg and pupal parasitoids can be related to the changes of the moth density in a delayed density-dependent manner. Furthermore, egg and pupal parasitoids cannot be overlooked as causal factors for the population cycles of forest Lepidoptera in general.     

Host locating behaviour of Leschenaultia exul and Patelloa pachypyga: two tachinid parasitoids of the forest tent caterpillar, Malacosoma disstria

Leschenaultia exul (Townsend) and Patelloa pachypyga (Aldrich & Webber) (Diptera: Tachinidae) are the principal larval parasitoids of the forest tent caterpillar (FTC) Malacosoma disstria (Hübner) (Lepidoptera: Lasiocampidae) in Canada. The response of these two fly species to M. disstria differs depending on the tree species on which the host feeds. In wind tunnel experiments, L. exul spent more time on the side of the tunnel containing volatiles from FTC frass and was attracted to the FTC-aspen poplar (Populus tremuloides Michx.) complex preferentially to the FTC-balsam poplar (Populus balsamifera L.) complex. Field bioassays confirmed that this fly species was preferentially attracted to the herbivore-aspen poplar complex as compared to the herbivore-balsam poplar complex. In field bioassays, P. pachypyga was also attracted preferentially to aspen poplar trees containing FTC larvae, compared to balsam poplar trees with host larvae.     

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.     

Possible physiological mechanisms for the differential susceptibility of two forest Lepidoptera to diflubenzuron

In an attempt to explain the physiological mechanisms for the differential susceptibility of insects to the chitin synthesis inhibitor, diflubenzuron, chitin content, chitin synthesis, and retention of ingested ¹⁴C-diflubenzuron in two forest Lepidoptera were investigated. The spruce budworm, Choristoneura fumiferana, a refractory species, had less chitin and retained less of the ingested material than the forest tent caterpillar, Malacosoma disstria, a species highly sensitive to diflubenzuron. No difference in the chitin synthesis pattern during the 6th stadium was observed in the two species. It is concluded that the primary reasons for the increased susceptibility of the forest tent caterpillar to this compound was the increased retention of ingested diflubenzuron and, to a lesser extent, the increased chitin content.     

Feeding by Hessian fly [Mayetiola destructor (Say)] larvae does not induce plant indirect defences

Abstract.  1. Recent research has addressed the function of herbivore-induced plant volatiles in attracting natural enemies of feeding herbivores. While many types of insect herbivory appear to elicit volatile responses, those triggered by gall insects have received little attention. Previous work indicates that at least one gall insect species induces changes in host-plant volatiles, but no other studies appear to have addressed whether gall insects trigger plant indirect defences.
2. The volatile responses of wheat to feeding by larvae of the Hessian fly Mayetiola destructor (Say) (Diptera: Cecidomyiidae) were studied to further explore indirect responses of plants to feeding by gall insects. This specialist gall midge species did not elicit a detectable volatile response from wheat plants, whereas a generalist caterpillar triggered volatile release. Moreover, Hessian fly feeding altered volatile responses to subsequent caterpillar herbivory.
3. These results suggest that Hessian fly larvae exert a degree of control over the defensive responses of their host plants and offer insight into plant-gall insect interactions. Also, the failure of Hessian fly larvae to elicit an indirect defensive response from their host plants may help explain why natural enemies, which often rely on induced volatile cues, fail to inflict significant mortality on M. destructor populations in the field.     

What causes population cycles of forest Lepidoptera?

Hypotheses for the causes of regular cycles in populations of forest Lepidoptera have invoked pathogen-insect or foliage-insect interactions. However, the available data suggest that forest caterpillar cycles are more likely to be the result of interactions with insect parasitoids, an old argument that seems to have been neglected in recent years.     

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