Winter Moth Research Articles

Detection and characterisation of three novel species of reovirus (Reoviridae), isolated from geographically separate populations of the winter moth Operophtera brumata (Lepidoptera: Geometridae) on Orkney

Geographically separate populations of winter moth ( Operophtera brumata L.) were sampled in heather habitats on the Orkney Isles in order to investigate the prevalence of virus pathogens. Reoviruses were isolated in 11 of the 13 winter moth populations sampled, with 3 novel species being detected. Two species of Cypoviridae (CPV) were isolated, Operophtera brumata CPV18 and O. brumata CPV19, with one host population suffering 46% infection prevalence of OpbuCPV19. A third virus, O. brumata Reovirus (OpbuRV), was isolated from both winter moth and a hymenopteran parasitoid wasp, Phobocampe tempestiva, which is abundant in these populations. This was identified as a non-occluded reovirus, which was clearly able to infect and persist in both the lepidopteran and the hymenopteran host. The genomes of the three viruses were characterised using gel electrophoresis and the virus structure was investigated using transmission electron microscopy. The relationship of these viruses with a baculovirus that also infects winter moth, OpbuNPV, was investigated, as well as the association of OpbuRV with P. tempestiva. The detection of such viruses is discussed with reference to studies of similar viruses in other lepidopteran and hymenopteran host systems.

Are polyphagous geometrid moths with flightless females adapted to budburst phenology of local host species?

The majority of studies demonstrating local adaptation of insect herbivores involve sessile species, particularly those with a parthenogentic phase to their life history or endophagous “parasites” of plants. Current arguments suggest the strength of selection determines whether local adaptation can or cannot take place. Therefore local adaptation should not be limited to species with such traits. We studied the ability of three polyphagous geometrid moths with flightless adult females (Erannisdefoliaria, Operophtera brumata and O. fagata) to synchronise their egg hatching with the budburst of a local host species in north east Scotland. A strong selection for hatching time is expected among generalist moths given the large variation in budburst phenology and an inability to hatch in synchrony with budburst decreases moth fitness substantially. In two successive seasons, we trapped emerging females from patches of five host species and recorded the temperature sum needed for 50% egg hatch of each brood laid by the trapped females. The hatching times of broods were compared against the average budburst time of the maternal host species in the study area. In addition, the trapping dates of each female were recorded. Only O. brumata showed synchrony with egg hatch and budburst which suggests local phenological adaptation to different host species. This could be maintained by selection and partial reproductive isolation between populations dwelling on different host species. No phenological adaptation was found in the other common geometrids of the study area.

Tree defoliation by winter moth Operophtera brumata L. during an outbreak affected by structure of forest landscape

Tree defoliation by winter moth Operophtera brumata L. (Geometridae, Lepidoptera) caterpillars was quantified in the Białowieża Forest (E Poland) during the recent outbreak (2002). To assess effects of forestry management on defoliation, comparisons were made between mixed species ( Tilio– Carpinetum) old-growth stands of natural origin in strictly protected primeval and managed fragments. This forest type was continuous in the primeval part, but highly fragmented in the managed part. The old-growth remnants in the latter part were usually surrounded by younger tree stands (often coniferous plantations). We predicted the winter moth densities would be reduced in fragmented stands due to dispersal mortality incurred by the early stage larvae. As predicted, all species of host trees suffered greater defoliation in the primeval forest. Hornbeam Carpinus betulus L. was the heaviest defoliated host. Other trees, which developed leaves synchronously with hornbeam – maple Acer platanoides L., lime Tilia cordata Mill., and pedunculate oak Quercus robur L. – were strongly affected as well. While we do not show a causal mechanism, greater dispersal mortality in fragmented stands is a most plausible explanation for the observed patterns of defoliation.

Spatial processes can determine the relationship between prey encounter rate and prey density

Theoretical models frequently assume that the rate at which a searching predator encounters prey increases linearly with prey density. In a recent experiment using great tits searching for winter moth caterpillars, the time to find the first prey item did not decline as quickly with density as the standard theory assumes. Using a spatial simulation model, we show that prey aggregation and/or spatially correlated searching behaviour by the predator can generate a range of relationships, including results that are qualitatively similar to those found in the great tit experiment. We suggest that further experiments are required to determine whether the explanation proposed here is correct, and that theoretical work is needed to determine how this behaviour is likely to influence the ecological and evolutionary dynamics of predator-prey communities.

Predatory Hypogaeic Beetles are Attracted to Buried Winter Moth (Lepidoptera: Geometridae) Pupae: Evidence Using a New Trap Design

A field study was carried out to determine whether small hypogaeic beetles prey on winter moth (Operophtera brumata [L.]) pupae. Small invertebrates were responsible for ca. 28% of pupal mortality at sites in British Columbia in 1991. Using a new trap design, adult Staphylinidae and beetle larvae were shown to actively search for buried pupae as indicated by significantly higher catches in hypogaeic traps baited with pupae than in unbaited, control traps. Baited and unbaited pitfalls were largely ineffective at capturing predatory larvae when compared to the baited hypogaeic traps. Many previous studies of predation on the soil-dwelling stages of insect herbivores may be biased towards the epigaeic component of the predator assemblage because they use pitfall traps alone. A combination of traps that includes hypogaeic traps should be considered for future studies.

Do multitrophic interactions override N fertilization effects on Operophtera larvae?

We examined how performance of Operophtera brumata (Lepidoptera) larvae was affected by nitrogen (N) fertilization of boreal forest understorey vegetation. We monitored larval densities on Vaccinium myrtillus plants for a period of 7 years in a field experiment. Preliminary results indicated that the N effect on larval densities was weak. To examine if this was due to indirect interactions with a plant pathogen, Valdensia heterodoxa, that share the same host plant, or due to top-down effects of predation, we performed both a laboratory feeding experiment (individual level) and a bird exclusion experiment (population level) in the field. At the individual level, altered food plant quality (changes in plant concentration of carbon, N, phenolics, or condensed tannins) due to repeated infection by the pathogen had no effect on larval performance, but both survival to the adult stage and adult weight were positively affected by N fertilization. Exclusion of insectivorous birds increased the frequency of larval damage on V. myrtillus shoots, indicating higher larval densities. This effect was stronger in fertilized than in unfertilized plots, indicating higher bird predation in fertilized plots. Predation may thus explain the lack of fertilization effect on larval densities in the field experiment. Our results suggest that top-down effects are more important for larval densities than bottom-up effects, and that bird predation may play an important role in population regulation of O. brumata in boreal forests.

The role of food plant and pathogen-induced behaviour in the persistence of a nucleopolyhedrovirus

Insect baculoviruses can survive between epidemics as infectious particles external to the host. Many pathogens persist in reservoirs, i.e., microhabitats where survival is enhanced, for example due to protection from the degrading effects of UV irradiation. However, the probability of infecting new susceptible hosts is usually reduced. Persistence of pathogens and their movement in and out of reservoirs is an important, albeit little understood, aspect of insect pathogen ecology. This study investigated interactions between the behaviour of infected insect hosts, virus distribution and plant species on the persistence of the winter moth ( Operophtera brumata) nucleopolyhedrovirus. Habitat influenced the persistence of infectious baculovirus in the field: virus on Sitka spruce ( Picea sitchensis) and oak ( Quercus robur) in forested areas retained more infectivity than virus on heather ( Calluna vulgaris) in an unshaded habitat. Plant species per se did not directly affect the persistence of virus on the foliage of potted seedlings. Virally infected insects had altered behaviour and moved down plants relative to control insects, whereas in other systems larvae show height-seeking behaviour. Consequently, the majority of virus particles were distributed on plant stems. In two experiments (one using winter moth NPV and one Mamestra brassicae NPV) virus persisted better on plant stems relative to foliage. Neonate larvae were shown to be able to acquire infections from tree stems contaminated with a low level of virus. These data suggest that plant stems may be important reservoirs for between-year persistence of this pathogen. The observed virus-induced changes in host behaviour in winter moth could enhance the viral persistence by increasing the deposition of occlusion bodies in these reservoirs.

Do sub‐Arctic winter moth populations in coastal birch forest exhibit spatially synchronous dynamics?

Summary 1 Populations of cyclic forest insects are known to exhibit large-scale spatially synchronous dynamics. Region-wide climatic disturbances, the Moran effect, have often been invoked as the mechanism underlying this synchrony. However, no previous study has been designed specifically to examine spatial population dynamics more closely and to evaluate underlying mechanisms. 2 We perform a study on winter moth populations in sub-Arctic birch forest, where habitat discontinuity was included in the study design, to evaluate if dispersal barriers would act to decouple dynamics. All the study populations were situated in the same climatic region where 10-year population cycles have been known to prevail. The total spatial extent of the study was smaller than the spatial scale of synchronous dynamics reported from previous studies on cyclic insects. 3 We found that moth populations had mainly synchronous dynamics within sites consisting of 1·8 km transects in prime moth habitats (mature birch forest) and within a large island with continuous birch forest. However, sites on different islands could be maximally out of phase. 4 These results suggest that climate was not synchronizing winter moth populations in this ecosystem and that biotic mechanisms are probably involved. 5 Our study highlights the value of performing studies with a particularly dedicated design for elucidating the patterns and mechanisms of spatio-temporal population dynamics.

Genetically variable nucleopolyhedroviruses isolated from spatially separate populations of the winter moth Operophtera brumata (Lepidoptera: Geometridae) in Orkney

Here we report a lepidopteran system in which a pathogen is both abundant and genotypically variable. Geographically separate populations of winter moth ( Operophtera brumata L.) were sampled in heather habitats on the Orkney Isles to investigate the prevalence of a pathogen, O. brumata Nucleopolyhedrovirus (OpbuNPV), within the natural system. Virus was recorded in 11 of the 13 winter moth populations sampled, with two populations suffering mortality due to virus at levels of ⩾50%. The virus genome from 200 single insect isolations was investigated for variation using restriction endonuclease digests. Twenty-six variants of OpbuNPV were detected using SalI. The polyhedrin gene of the virus was partially sequenced, allowing the relationship between the 26 variants to be portrayed as a cladogram. The phylogenetic relationship between OpbuNPV and other known baculovirus polyhedrin gene sequences was also established. The discovery of virus at such high prevalence is discussed with reference to occurrence and genetic variation of pathogens in other lepidopteran host populations. This study shows encouraging results for further studies into the role of pathogens in the regulation of host insect populations.

Oviposition preference of Anthocoris nemorum and A. nemoralis for apple and pear

Abstract Anthocoris nemorum L. and Anthocoris nemoralis Fabricius (Heteroptera: Anthocoridae) are important predators of insect pests in pome fruit. Females insert their eggs in leaf tissue. The females’ choice of oviposition site is important for the subsequent distribution of nymphs on host plants. Oviposition preference for apple and pear leaves was tested in the laboratory in four experiments (experiments 1–4). In three experiments it was tested whether simulated insect damage to leaves (experiments 5 and 6) or the presence of prey (experiment 7) influenced oviposition preference. The effect of the presence of prey was only tested for A. nemorum on apple leaves. There was a highly significant anthocorid species × plant interaction for the number of eggs laid on apple and pear leaves. Anthocoris nemorum laid more eggs on apple than on pear leaves, while A. nemoralis preferred pear. Anthocoris nemorum's preference for apple increased over the 6‐week period in which experiments 1–4 were performed, from 66% to 91% eggs laid on apple leaves. No change over time in preference was found for A. nemoralis. Across experiments 1–4, the majority of A. nemorum eggs were laid near leaf margins, whereas eggs of A. nemoralis were more commonly found in the leaf centre, 5 mm or more from the margin, with a highly significant leaf region × species interaction. There was no significant difference in preference for leaf side between A. nemorum and A. nemoralis, but there was a highly significant plant × leaf side × experiment interaction. Thus, more eggs were laid on the ventral than on the dorsal side of pear leaves in experiment 4, while significantly more eggs were laid on the dorsal side of apple leaves in experiments 3 and 4. Choice tests between damaged and healthy leaves showed that A. nemorum laid significantly more eggs on the damaged leaves, while A. nemoralis preferred healthy leaves. Anthocoris nemorum showed a near‐significant preference for ovipositing on leaves with eggs of Operophtera brumata (Lepidoptera: Geometridae). The oviposition preferences found correspond to the natural distribution of these predators in apple and pear orchards. The preference of A. nemorum for leaf margins, and of A. nemoralis for the leaf centre as an oviposition site, supports earlier observations. A preference for leaf side for oviposition site has not been reported earlier. Preference for damaged leaves could help A. nemorum to locate prey in a field situation.

Identification of polyenic hydrocarbons from the northern winter moth, Operophtera fagata , and development of a species specific lure for pheromone traps

In order to elucidate the composition of the female sex pheromone of the northern (beech) winter moth, Operophtera fagata Scharf. (Lepidoptera: Geometridae), ovipositor extracts of unmated, calling females were analysed by gas chromatography with simultaneous electroantennographic and flame ionization detection (GC-EAD/FID). Male antennal responses indicated three active components, two of which had distinct matching peaks in the FID trace. Using coupled gas chromatography- mass spectrometry (GC-MS), these two compounds were identified as (9Z)-nonadecene (9Z-19:Hy), and (6Z,9Z)-nonadecadiene (6Z9Z-19:Hy), respectively. The third component, present in very small amounts only, was identified as (1,3Z,6Z,9Z)-nonadecatetraene (1,3Z6Z9Z-19:Hy), known as the sex pheromone of the common winter moth, O. brumata. Field tests revealed that traps baited with 6Z9Z-19:Hy and 1,3Z6Z9Z-19:Hy caugth large numbers of male O. fagata. Both compounds were found to be essential for attraction of O. fagata. In addition, the diene prevented captures of co-occurring O. brumata. In contrast, 9Z-19:Hy neither influenced the attractiveness of the two-component mixture towards O. fagata nor contributed to bait specificity. A binary mixture of 6Z9Z-19:Hy and 1,3Z6Z9Z-19:Hy in a ratio of 10:1, applied to pieces of rubber tubing, constituted a highly attractive and species-specific bait for O. fagata, which can be used for monitoring of the flight of this defoliator pest of deciduous forests.

Central assumptions of predator–prey models fail in a semi–natural experimental system

The relationship between the encounter rate of predators with prey and the density of this prey is fundamental to models of predator-prey interactions. The relationship determines, among other variables, the rate at which prey patches are depleted, and hence the impact of predator populations on their prey, and the optimal spatial distribution of foraging effort. Two central assumptions that are made in many models are that encounter rate is directly proportional to prey density and that it is independent of the proportion of prey already removed, other than via the decreased density. We show here, using captive great tits searching for winter moth caterpillars in their natural hiding positions, that neither of these assumptions hold. Encounter rate increased less than directly in proportion to prey density, and it depended not only on the current density of prey, but also on the proportion of prey already removed by previous foragers. Both of these effects are likely to have major consequences for the outcome of predator-prey interactions.

Interactions between predatory ground beetles, the winter moth and an introduced parasitoid on the Lower Mainland of British Columbia

This study describes assemblages of predatory ground beetles in native birch forest and derived blueberry farms on the Lower Mainland of British Columbia and investigates their role in the successful biological control of winter moth ( Operopthera brumata L.) by the parasitoid Cyzenis albicans Fallén. Pitfall traps at four sites in 1991 and 1992 indicated that natural birch forest was dominated by native carabid species, while open blueberry was dominated by introduced species and some native xerophilous species. Predation by generalist invertebrates was an important mortality factor for both winter moth and C. albicans at all sites and was greatest in the first 2 weeks after pupation. Ground beetles including Harpalus affinus (Schrank) and Pterostichus spp. consumed winter moth pupae in arenas but adult beetles rarely consumed C. albicans pupae. Much of the overall predation was due to smaller invertebrates including beetle larvae. Predators actively searched for buried pupae and selected among pupal types based on condition (dead, parasitised or healthy) and size. Prey-selection differed among sites but C. albicans pupae were preferred at three of the four sites. At the remaining site, an organic blueberry farm, over 90% of both parasitised and healthy pupae were consumed. High levels of predation did not affect the levels of parasitism by C. albicans at the sites. This herbivore–parasitoid–predator system demonstrates the importance of naturally occurring predatory invertebrates for successful integrated pest management, even where they prey upon a biological control agent.

Density‐dependent melanism in sub‐arctic populations of winter moth larvae (Operophtera brumata)

Abstract. 1. The aim of this 4‐year observational study was to test for the presence of direct and delayed density‐dependent larval melanism in the geometrid moth species Operophtera brumata (winter moth) in northern Norway.2. Data from many populations with a wide range of population densities in time and space facilitated statistical analyses that could separate the effects of current and past density. The data also included different phases of the 10‐year population cycle of this species so that eventual non‐linear density effects due to population phase could be detected.3. The results showed that the prevalence of melanism had a strong positive, linear relation to population density within years, whereas there was no evidence for a delayed effect from the year before or dependency on the phase of the population cycle.4. In combination, these results limit the range of possible explanations of larval melanism in this outbreaking species. The possible reasons why winter moth larvae might benefit from crowding‐induced melanism are discussed.

Host shifting by Operophtera brumata into novel environments leads to population differentiation in life‐history traits

Abstract. 1. Operophtera brumata L. (Lepidoptera: Geometridae), a polyphagous herbivore usually associated with deciduous trees such as oak Quercus robur L., has expanded its host range to include the evergreen species heather Calluna vulgaris (L.) Hull and, most recently, Sitka spruce Picea sitchensis (Bong.) Carrière.2. Phenology, morphology, and survival of O. brumata were measured at several life‐history stages in populations from the three different host plant communities sampled from a range of geographical locations. The data were used to test for population differences, reflecting the marked differences in host‐plant secondary chemistry, growth form, and site factors such as climate. The hypothesis that spruce‐feeding populations originated from populations feeding on moorland, commonly sites of coniferous afforestation, was also tested.3. Altitude, not host plant species, was the major influence on the timing of adult emergence. An effect of insect population independent of altitude was found, implying that additional unidentified factors contribute to this phenological variation. Larval survival and adult size varied between populations reared on different host plant species. Survival of larvae was affected negatively when reared on the novel host plant, Sitka spruce, versus the natal plant (oak or heather) but oak and heather‐sourced insects did not differ in survivorship on Sitka spruce.4. Host range extension into novel environments has resulted in population differentiation to the local climate, demonstrating that host shifts pose challenges to the herbivore population greater than those offered by the host plant alone. The hypothesis that Sitka spruce feeding populations have arisen predominantly from moorland feeding populations was not supported.

Biocontrol Potential of Entomopathogenic Nematodes Against Winter Moths (Operophtera brumata and O. fagata) (Lepidoptera: Geometridae) Infesting Urban Trees

Infectivity and biocontrol potential of entomopathogenic nematodes against winter moths (Operophtera brumata and O. fagata)pupating in the soil were examined in laboratory, semi-field and field conditions. A pilot experiment conducted in the field showed that Steinernema feltiae was completely ineffective against pupae of these moths in the soil. Subsequent laboratory tests revealed that none of the tested species (i.e. S. feltiae, S. affinae, S. carpocapsae, Heterorhabditis megidis and H. bacteriophora) could colonise the pupae, while mature larvae descending to the soil for pupation and prepupae were highly susceptible to nematode infection. No differences were observed between O. brumata and O. fagata in susceptibility to nematodes. In laboratory experiments H. megidis applied at 1.5×10 <italic>5</italic> infective juveniles (IJ) m <italic>−2</italic> infected almost 100% of insects exposed for 6 days in the soil. It was significantly more infective than H. bacteriophora (73-77%) and Steinernema species (29-50%). H. megidis was also highly effective in semi-field conditions when applied at an even lower dose, i.e. 10 <italic>5</italic> IJ m <italic>−2</italic> . After a 45-day experiment, only 3% of insects descending for pupation survived in the soil pre-treated with this species. This was significantly less than in soil with S. feltiae (43%) and control treated with water only (59%). Very high efficacy of H. megidis and a relatively easy method for its field application through ground spraying gives some promise for environmentally safe and successful biological control of winter moths during their pupation in the soil. The low application rate required and recycling in the host could be additional advantages for economic and long lasting protection of high value trees, particularly those in urban parks and forests.

Phenological variation as protection against defoliating insects: the case of Quercus robur and Operophtera brumata.

Phenological synchrony between budburst and emergence of larvae is critical for the fitness of many spring-feeding insect herbivores. Therefore, large intraspecific variation in timing of budburst of the host may have a negative effect on the herbivore. We studied how asynchrony between emergence of larvae and budburst affects the fitness of Operophtera brumata (Lepidoptera: Geometridae), a major defoliator of Quercus robur, which can adapt to the phenology of a single tree. It is known that, in maturing leaves of Q. robur, accumulation of condensed tannins has a negative effect on growth of O. brumata. However, there is no information about the effect of hydrolysable tannins and other phenolics that are potential antifeedants. In this study, we also analysed changes in secondary chemistry of the foliage of Q. robur and how different compounds are correlated with growth and survival of O. brumata. The effect of asynchrony on O. brumata was studied in rearing experiments. The neonate larvae were incubated without food for different periods of time. The decline in nutritional quality of foliage was estimated by rearing cohorts of larvae with manipulated hatching times on the leaves of ten individual Q. robur trees. For the chemical analysis, the foliage of these trees was sampled at regular intervals. In the absence of foliage, mortality of neonate larvae started to increase exponentially soon after the larvae emerged. If the larvae missed budburst, the decline in nutritional quality of the foliage led to increased mortality and lower body mass (= fecundity). Hydrolysable tannins were not significantly correlated with performance of the larvae. Only condensed tannins were found to correlate negatively with the growth and survival of O. brumata. Certain individual trees were unsuitable hosts for O. brumata because the decline in quality of the foliage was very rapid. Based on regression equations for increasing rate of mortality and decreasing fecundity, we calculated that a relatively small mismatch of +/-30 degree days between budburst and hatching of larvae leads to a 50% decrease in the fitness of O. brumata. Thus, large phenological variation within a Q. robur stand can limit the colonisation of neighbouring trees by dispersing larvae. Furthermore, the hybridisation of moths adapted to phenologically different trees may lead to maladapted phenology of their offspring.

Genetic differentiation in the winter pine processionary moth (Thaumetopoea pityocampa--wilkinsoni complex), inferred by AFLP and mitochondrial DNA markers.

The winter pine processionary moth has become an important pine pest in the last century, as a consequence of the spread of pine cultivation in the Mediterranean region. The pattern of genetic differentiation of this group, that includes two sibling species (Thaumetopoea pityocampa and Th. wilkinsoni), has been studied in nine populations using amplified fragment length polymorphism (AFLP) and single strand conformation polymorphism-sequence analysis (SSCP) of the mitochondrial cytochrome oxidase 1 (COI) and cytochrome oxydase 2 (COII). Results indicate the existence of strong genetic differentiation between the two species that became separated before the Quaternary ice ages. Moreover data indicate that Th. pityocampa has a strong geographical structure, particularly evident at the nuclear level, where all pairwise phiST resulted to be highly significant and individuals from the same population resulted to be strongly clustered when an individual tree was reconstructed. The estimates of the absolute number of migrants between populations (Nm), obtained from mitochondrial and nuclear DNA markers, suggest that gene flow is low and that a gender-related dispersal could occur in this species. The males appear to disperse more than females, contributing to the genetic diversity of populations on a relatively wide range, reducing the risks of inbreeding and the genetic loss associated with bottlenecks occurring in isolated populations.

Host‐mediated effects of feeding by winter moth on the survival of Euceraphis betulae

Abstract 1. A potential host‐mediated interaction between the birch‐feeding aphid Euceraphis betulae Koch (Homoptera: Aphididae) and Operophtera brumata L. (Geometridae: Lepidoptera) was investigated, by measuring survival and feeding preference responses of E. betulae to leaves damaged recently by O. brumata larvae.2. Euceraphis betulae survival was considerably lower on damaged foliage than on undamaged foliage, both in field experiments using cages and on potted saplings in laboratory tests.3. Euceraphis betulae did not avoid damaged foliage in field experiments although they did avoid damaged foliage in laboratory preference tests using individual leaves. A link between aphid preference and performance on damaged foliage could therefore be demonstrated in the greenhouse/laboratory but not in the field. Multiple factors may influence feeding preference in the more complex field environment.4. These results suggest that host‐mediated competition may occur between O. brumata and E. betulae, especially in outbreak years when winter moth populations are very high.

Escape from pupal predation as a potential cause of outbreaks of the winter moth, Operophtera brumata

The winter moth, Operophtera brumata, shows varying population dynamics in different host plant habitats. Populations in Sitka spruce, Picea sitchensis, plantations and in Scottish moorlands have a tendency to outbreak that is not shown by winter moth in lowland oak woods. Since pupal predators have previously been identified as being important for the regulation of winter moth in a lowland oak wood it was hypothesized that invertebrate pupal predators were failing to control winter moth in outbreak populations. This hypothesis was tested by comparing the abundance of invertebrate predators and patterns of spatially density dependent pupal predation across habitats. Several results supported this hypothesis. Carabid predators of winter moth were one or two orders of magnitude more abundant in oak woods than in moorland or spruce habitats. Staphylinid predators were also more abundant in high winter moth density oak woods than in any other habitat. Beetle predation of tagged cocoons in the field was inversely density dependent in Highland moors in experiments in 1999 and 2000, and in Sitka spruce in 1999. However, in opposition to our hypothesis, pupal predation was also inversely density dependent in oak woods in 2000, although this result may be explained by the low range of winter moth densities in the field that year. These results are discussed in relation to the role of natural enemies in regulating winter moth populations and the differences in life‐history of the beetle predators in different habitats.