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Climate change increases the insect abundance, especially in Arctic ecosystems. Insect herbivory also significantly increases plant emissions of volatile organic compounds (VOCs), which are highly reactive in the atmosphere and play a crucial role in atmospheric chemistry and physics. However, it is unclear how the effects of insect herbivory on VOC emissions interact with climatic changes, such as warming and increased cloudiness. We assessed how experimental manipulations of temperature and light availability in subarctic tundra, that had been maintained for 30 years at the time of the measurements, affect the VOC emissions from a widespread dwarf birch (Betula nana) when subjected to herbivory by local geometrid moth larvae, the autumnal moth (Epirrita autumnata) and the winter moth (Operophtera brumata). Warming and insect herbivory on B. nana stimulated VOC emission rates and altered the VOC blend. The herbivory‐induced increase in sesquiterpene and homoterpene emissions were climate‐treatment‐dependent. Many herbivory‐associated VOCs were more strongly induced in the shading treatment than in other treatments. We showed generally enhanced tundra VOC emissions upon insect herbivory and synergistic effects on the emissions of some VOC groups in a changing climate, which can have positive feedbacks on cloud formation. Furthermore, the acclimation of plants to long‐term climate treatments affects VOC emissions and strongly interacts with plant responses to herbivory. Such acclimation complicates predictions of how climate change, together with interacting biotic stresses, affects VOC emissions in the high latitudes., Climate change increases the insect abundance, especially in arctic ecosystems. Insect herbivory also significantly increases plant emissions of volatile organic compounds (VOCs), which are highly reactive in the atmosphere and play a crucial role in atmospheric chemistry and physics. We showed generally enhanced tundra VOC emissions upon insect herbivory and synergistic effects on the emissions of some VOC groups in a changing climate, which can have positive feedbacks on cloud formation.

Key message Whereas cold temperatures and artificial smoke pollution (during World War II) cause negative pointer years in northern Fennoscandian downy birches, mass outbreaks of Epirrita autumnata L. and Operophtera brumata Bkh. are the strongest growth-influencing and -synchronizing factor. Abstract Variations in radial tree growth of downy birch can be affected by short-term environmental changes. Here we examine the tree-ring width of downy birch trees for signals caused by three different disturbance factors: climatic extremes, insect outbreaks, and artificial smoke employed to hide the German battleship Tirpitz during World War II at the Kåfjord in northern Norway. Besides growth/climate response analysis using gridded climate data and a tree ring width-chronology of the studied birch trees, we analyze missing rings, pointer years, and the ability of recovery after severe disturbances on an individual tree basis using the percent change of the trees’ basal area increment with respect to a reference period. The downy birches reveal (1) a significant positive correlation with May and June temperatures, (2) a high growth sensitivity to moth epidemics (Epirrita autumnata L. and Operophtera brumata Bkh.), and (3) a distinct growth deviation in the year of intense but short-term artificial smoke pollution. We conclude that downy birches are not exclusively sensitive to unusual cold temperatures, but short-term artificial pollution and insect outbreaks cause similar growth declines in the year of occurrence. Whereas for temperature and pollution, the growth response is less coherent among trees, and fades within 2 years, the insect outbreaks synchronize growth among all downy birches. Growth declines during years of mass insect outbreaks can further be amplified by cold May and June temperatures. Our findings thereby improve the understanding of competing disturbance factors on single birch trees but also on the growth of a whole site in Fennoscandia and facilitates the detection of disturbances in birch chronologies.

Despite active research, antiherbivore activity of specific plant phenolics remains largely unresolved. We constructed silver birch (Betula pendula) lines with modified phenolic metabolism to study the effects of foliar flavonoids and condensed tannins on consumption and growth of larvae of a generalist herbivore, the autumnal moth (Epirrita autumnata). We conducted a feeding experiment using birch lines in which expression of dihydroflavonol reductase (DFR), anthocyanidin synthase (ANS) or anthocyanidin reductase (ANR) had been decreased by RNA interference. Modification-specific effects on plant phenolics, nutrients and phenotype, and on larval consumption and growth were analyzed using uni- and multivariate methods. Inhibiting DFR expression increased the concentration of flavonoids at the expense of condensed tannins, and silencing DFR and ANR decreased leaf and plant size. E. autumnata larvae consumed on average 82% less of DFRi plants than of unmodified controls, suggesting that flavonoids or glandular trichomes deter larval feeding. However, larval growth efficiency was highest on low-tannin DFRi plants, indicating that condensed tannins (or their monomers) are physiologically more harmful than non-tannin flavonoids for E. autumnata larvae. Our results show that genetic manipulation of the flavonoid pathway in plants can effectively be used to produce altered phenolic profiles required for elucidating the roles of low-molecular weight phenolics and condensed tannins in plant–herbivore relationships, and suggest that phenolic secondary metabolites participate in regulation of plant growth.

With warming climate many species are predicted to shift their distributions toward the poles. However, climate change models developed to predict species distributions do not always incorporate interactions between them. The northerly shift of the boreal forest and associated dwarf shrub communities will be directly affected by warming. But warming will also indirectly affect plant communities via impacts on the intensity and frequency of associated insect outbreaks. We present a general model exploring plant host herbivory in response to the balance between insect crowding, host consumption and climate. We examined how these factors dictate the feeding preference of Epirrita autumnata larvae during an outbreak on dwarf shrub vegetation in Sub-arctic Fennoscandia. Data were collected from an outdoor experiment investigating future climate change scenarios (elevated CO2 and temperature) on the dwarf shrub community that included deciduous (Vaccinium myrtillus) and evergreen species (V. vitis-idaea and Empetrum nigrum). We observed that larval crowding was independent of treatment under outbreak conditions. We also tested and confirmed model predictions that larvae would prefer monospecific stands of either deciduous shrubs or its evergreen competitors. For current climate conditions, larvae had a preference to consume more deciduous shrubs in mixed stands. However, at elevated temperature bilberry consumption and herbivore pressure was lower, particularly in mixed stands. Our results show that during future warming, E. autumnata herbivory could promote the success of thermophile deciduous species and possible northward migration. Insect behaviour and preferences should therefore be considered when predicting future vegetation movements responding to warming.

The following article, Bokhorst, S., Berg, M.P., Edvinsen, G.K., Ellers, J., Heitman, A., Jaakola, L., ... Bjerke, J.W. (2018). Impact of Multiple Ecological Stressors on a Sub-Arctic Ecosystem: No Interaction Between Extreme Winter Warming Events, Nitrogen Addition and Grazing. Frontiers in Plant Science, 9, can be accessed at https://doi.org/10.3389/fpls.2018.01787. Climate change is one of many ongoing human-induced environmental changes, but few studies consider interactive effects between multiple anthropogenic disturbances. In coastal sub-arctic heathland, we quantified the impact of a factorial design simulating extreme winter warming (WW) events (7 days at 6–7∘C) combined with episodic summer nitrogen (+N) depositions (5 kg N ha-1) on plant winter physiology, plant community composition and ecosystem CO2 fluxes of an Empetrum nigrum dominated heathland during 3 consecutive years in northern Norway. We expected that the +N would exacerbate any stress effects caused by the WW treatment. During WW events, ecosystem respiration doubled, leaf respiration declined (-58%), efficiency of Photosystem II (Fv/Fm) increased (between 26 and 88%), while cell membrane fatty acids showed strong compositional changes as a result of the warming and freezing. In particular, longer fatty acid chains increased as a result of WW events, and eicosadienoic acid (C20:2) was lower when plants were exposed to the combination of WW and +N. A larval outbreak of geometrid moths (Epirrita autumnata and Operophtera brumata) following the first WW led to a near-complete leaf defoliation of the dominant dwarf shrubs E. nigrum (-87%) and Vaccinium myrtillus (-81%) across all experimental plots. Leaf emergence timing, plant biomass or composition, NDVI and growing season ecosystem CO2 fluxes were unresponsive to the WW and +N treatments. The limited plant community response reflected the relative mild winter freezing temperatures (-6.6∘C to -11.8∘C) recorded after the WW events, and that the grazing pressure probably overshadowed any potential treatment effects. The grazing pressure and WW both induce damage to the evergreen shrubs and their combination should therefore be even stronger. In addition, +N could have exacerbated the impact of both extreme events, but the ecosystem responses did not support this. Therefore, our results indicate that these sub-arctic Empetrum-dominated ecosystems are highly resilient and that their responses may be limited to the event with the strongest impact.

International audience; We explore a common feature of insect population dynamics, interspecific synchrony, which refers to synchrony in population dynamics among sympatric populations of different species. Such synchrony can arise via several possible mechanisms, including shared environmental effects and shared trophic interactions, but distinguishing the relative importance among different mechanisms can be challenging. We analyze interannual time series of population densities of the larch budmoth,Zeiraphera griseana(Lepidoptera: Tortricidae), along with six sympatric larch-feeding folivores from a site in the European Alps 1952-1979. These species include five lepidopterans,Exapate duratella,Ptycholomoides aeriferana,Spilonota laricana,Epirrita autumnataandTeleiodes saltuum, and one hymenopteran sawflyPristiphora laricis. We document that the highly regular oscillatory behavior (period 9-10 years) ofZ.griseanapopulations is similarly evident in the dynamics of most of the sympatric folivores. We also find that all of the sympatric species are phase synchronized withZ.griseanapopulations with half of the sympatric species exhibiting nonlagged phase synchrony and three of the species exhibiting 2-5 year lags behindZ.griseanapopulations. We adapt a previously developed tritrophic model ofZ.griseanadynamics to explore possible mechanisms responsible for observed phase synchronization. Results suggest that either shared stochastic influences (e.g., weather) or shared parasitoid impacts are likely causes of nonlagged phase synchronization. The model further indicates that observed patterns of lagged phase synchronization are most likely caused by either shared delayed induced host plant defenses or direct density-dependent effects shared withZ.griseana.

Insect herbivory is known to augment emissions of biogenic volatile organic compounds (BVOCs). Yet few studies have quantified BVOC responses to insect herbivory in natural populations in pan-Arctic regions. Here, we assess how quantitative and qualitative BVOC emissions change with increasing herbivore feeding intensity in the Subarctic mountain birch (Betula pubescens var pumila (L.)) forest. We conducted three field experiments in which we manipulated the larval density of geometrid moths (Operophtera brumata and Epirrita autumnata), on branches of mountain birch and measured BVOC emissions using the branch enclosure method and gas chromatography-mass spectrometry. Our study showed that herbivory significantly increased BVOC emissions from the branches damaged by larvae. BVOC emissions increased due to insect herbivory at relatively low larvae densities, causing up to 10% of leaf area loss. Insect herbivory also changed the blend composition of BVOCs, with damaged plants producing less intercorrelated BVOC blends than undamaged ones. Our results provide a quantitative understanding of the relationship between the severity of insect herbivore damage and emissions of BVOCs at larvae densities corresponding to background herbivory levels in the Subarctic mountain birch. The results have important and practical implications for modeling induced and constitutive BVOC emissions and their feedbacks to atmospheric chemistry. arctic, biotic stress, geometrid moth, insect herbivory, mountain birch, stress severity, volatile organic compounds

1. Population dynamics and interactions that vary over a species' range are of particular importance in the context of latitudinal clines in biological diversity. Winter moth (Operophtera brumata) and autumnal moth (Epirrita autumnata) are two species of eruptive geometrids that vary widely in outbreak tendency over their range, which generally increases from south to north and with elevation. 2. The predation pressure on geometrid larvae and pupae over an elevational gradient was tested. The effects of background larval density and bird occupancy of monitoring nest boxes on predation rates were also tested. Predation on larvae was tested through exclusion treatments at 20 replicate stations over four elevations at one site, while pupae were set out to measure predation at two elevations at three sites. 3. Larval densities were reduced by bird predation at three lower elevations, but not at the highest elevation, and predation rates were 1.9 times higher at the lowest elevation than at the highest elevation. The rate of predation on larvae was not related to background larval density or nest box occupancy, although there were more eggs and chicks at the lowest elevation. There were no consistent differences in predation on pupae by elevation. 4. These results suggest that elevational variation in avian predation pressure on larvae may help to drive elevational differences in outbreak tendency, and that birds may play a more important role in geometrid population dynamics than the focus on invertebrate and soil predators of previous work would suggest.

We studied the feeding ecology of the common frog Rana temporaria in a population located at the subarctic taiga-tundra frontier in Finland over two consecutive years (2002, 2003). In the summer of 2003, an unexpected gradation of autumnal moth (Epirrita autumnata) larvae occurred at our study site. This situation allowed us to study the changes in the diet composition of the same common frog population between a presumably average and an extraordinary year with respect to the gradation of one of the prey species. We found clear evidence for the opportunistic feeding behaviour of common frogs in a natural population, as reflected by the mass appearance of Epirrita autumnata larvae in the frogs’ stomach content in 2003. Further, our results draw attention to the fact that a one-year study on the feeding ecology of an opportunistic species can result in flawed conclusions.

Tree mortality from insect infestations can significantly reduce carbon storage in forest soils. In subarctic birch forests (Betula pubescens), ecosystem C cycling is largely affected by recurrent outbreaks of defoliating geometrid moths (Epirrita autumnata, Operophtera brumata). Here, we show that soil C stocks in birch forests across Fennoscandia did not change up to 8 years after moth outbreaks. We found that a decrease in woody fine roots was accompanied by a lower soil CO2 efflux rate and a higher soil N availability following moth outbreaks. We suggest that a high N availability and less ectomycorrhiza likely contributed to lowered heterotrophic respiration and soil enzymatic activity. Based on proxies for decomposition (heterotrophic respiration, phenol oxidase potential activity), we conclude that a decrease in decomposition is a prime cause why soil C stocks of mountain birch forest ecosystems have not changed after moth outbreaks. Compared to disturbed temperate and boreal forests, a CO2-related positive feedback of forest disturbance on climate change might therefore be smaller in subarctic regions. Betula pubescens; disturbed subarctic forests; Epirrita autumnata; heterotrophic soil respiration; Operophtera brumata; root biomass; soil carbon sequestration; soil CO2 efflux; soil enzyme activity; structural equation modelling.

Sub-arctic birch forests (Betula pubescens Ehrh. ssp. czerepanovii) periodically suffer large-scale defoliation events caused by the caterpillars of the geometrid moths Epirrita autumnata and Operophtera brumata. Despite their obvious influence on ecosystem primary productivity, little is known about how the associated reduction in belowground C allocation affects soil processes. We quantified the soil response following a natural defoliation event in sub-arctic Sweden by measuring soil respiration, nitrogen availability and ectomycorrhizal fungi (EMF) hyphal production and root tip community composition. There was a reduction in soil respiration and an accumulation of soil inorganic N in defoliated plots, symptomatic of a slowdown of soil processes. This coincided with a reduction of EMF hyphal production and a shift in the EMF community to lower autotrophic C-demanding lineages (for example, /russula-lactarius). We show that microbial and nutrient cycling processes shift to a slower, less C-demanding state in response to canopy defoliation. We speculate that, amongst other factors, a reduction in the potential of EMF biomass to immobilise excess mineral nitrogen resulted in its build-up in the soil. These defoliation events are becoming more geographically widespread with climate warming, and could result in a fundamental shift in sub-arctic ecosystem processes and properties. EMF fungi may be important in mediating the response of soil cycles to defoliation and their role merits further investigation. Electronic supplementary material The online version of this article (doi:10.1007/s10021-016-0026-7) contains supplementary material, which is available to authorized users.

Climate change may facilitate shifts in the ranges and the spread of insect pests, but a warming climate may also affect herbivorous insects adversely if it disrupts the locally adapted synchrony between the phenology of insects and that of their host plant. The ability of a pest species to colonize new areas depends on its ability to adjust the timing of phenological events in its life cycle, particularly at high latitudes where there is marked seasonality in temperature and day length. Here we incubated eggs of three species of geometrid moth, Epirrita autumnata, Operophtera brumata and Erannis defoliaria from different geographical populations (E. autumnata and O. brumata from Northern Finland, E. autumnata and E. defoliaria from Southern Finland and all three species from Germany) in a climate chamber at a constant temperature to determine the relative importance of geographic origin in the timing of egg hatch measured in terms of cumulative temperature sums (degree days above 5°C, DD5); i.e. the relative importance of local adaptation versus phenotypic plasticity in the timing of egg hatch. In all three species, eggs from northern populations required a significantly lower temperature sum for hatching than eggs from southern populations, but the differences between them in temperature sum requirements varied considerably among species, with the differences being largest for the earliest hatching and northernmost species, E. autumnata, and smallest for the southern, late-hatching E. defoliaria. In addition, the difference in hatch timing between the E. autumnata eggs from Southern Finland and Germany was many times greater than the difference between the two Finnish populations of E. autumnata, despite the fact that the geographical distances between these populations is similar. We discuss how these differences in hatching time may be explained by the differences in hatch-budburst synchrony and its importance for different moth species and populations. We also briefly reflect on the significance of photoperiod, which is not affected by climate change. It is a controller that works parallel or in addition to temperature sum both for egg hatch in moths and bud burst of their host plants.

A seven-year long, two-factorial experiment using elevated temperatures (5 °C) and CO2 (concentration doubled compared to ambient conditions) designed to test the effects of global climate change on plant community composition was set up in a Subarctic ecosystem in northernmost Sweden. Using point-frequency analyses in permanent plots, an increased abundance of the deciduous Vaccinium myrtillus, the evergreens V. vitis-idaea and Empetrum nigrum ssp. hermaphroditum and the grass Avenella flexuosa was found in plots with elevated temperatures. We also observed a possibly transient community shift in the warmed plots, from the vegetation being dominated by the deciduous V. myrtillus to the evergreen V. vitis-idaea. This happened as a combined effect of V. myrtillus being heavily grazed during two events of herbivore attack—one vole outbreak (Clethrionomys rufocanus) followed by a more severe moth (Epirrita autumnata) outbreak that lasted for two growing seasons—producing a window of opportunity for V. vitis-idaea to utilize the extra light available as the abundance of V. myrtillus decreased, while at the same time benefitting from the increased growth in the warmed plots. Even though the effect of the herbivore attacks did not differ between treatments they may have obscured any additional treatment effects. This long-term study highlights that also the effects of stochastic herbivory events need to be accounted for when predicting future plant community changes.

Ericaceous dwarf shrubs, such as bilberry, Vaccinium myrtillus, have an important role in nutrient cycling of boreal forests, but in metal polluted environments they also form a link between heavy metal pool of the soil, primary consumers and upper trophic levels. From the viewpoint of metal transfer in a food chain, we document metallic element (As, Ca, Cd, Co, Cu, Mn, Mo, Ni, Pb, Se, Zn) concentrations in leaves, berries and herbivorous larvae of V. myrtillus around a Finnish copper-nickel smelter and compare those with levels in relatively unpolluted reference sites, and with levels documented in soil and feces (a proxy of dietary levels) of an insectivorous bird, the pied flycatcher, Ficedula hypoleuca. Herbivorous larvae of the autumnal moth, Epirrita autumnata (Lepidoptera: Geometridae), grown experimentally on V. myrtillus, showed slower growth rate but not higher mortality in the polluted area. In general, metal levels in leaves, berries and larvae were higher in the polluted area and comparable to those reported at other smelter sites in Europe. The levels of the main toxic metals (As, Cd, Cu, Ni, Pb) followed the general pattern: soil > bird feces > leaves > larvae = berries, and levels in V. myrtillus, E. autumnata and F. hypoleuca reflected soil metal levels. The lowest levels were found in those matrices that are most important sources of food for birds and humans, i.e. leaf-eating larvae and berries, reducing a risk of toxic effects.

‘Cry for help’ hypothesis predicts that attraction of predators with chemical or visual cues can decrease insect damage of plants. Visual cues involve changes in photosynthetic activity and the reflectance of leaves, and there is some evidence that birds may use these changes as foraging cues. However, changes in the visual properties of leaves have not been quantified and it is not known how birds see these changes. We also presented and tested a new ‘reduction in camouflage’ hypothesis (not mutually exclusive with cry for help) stating that herbivore-mediated changes in leaf colour can increase the conspicuousness of herbivore against leaves. To define changes in the visual properties of leaves, their detectability to birds, and whether these changes affect the conspicuousness of herbivore, we manipulated the level of herbivory in silver birch trees (Betula pendula) with autumnal moth (Epirrita autumnata) larvae, and used blue tit (Cyanistes caeruleus) vision models to images of leaves and larvae. Hue, luminance (lightness), contrast, light transmission, chlorophyll content, photosynthetic activity and water content of the leaves were compared between herbivore-damaged and control trees. The leaves of herbivore-damaged trees had a decreased chlorophyll a concentration, increased contrast and they reflected more longer wavelengths. However, these changes are likely not obvious to birds. In contrast to our expectation, there were only minor differences in conspicuousness of larvae against the leaves of damaged trees, which may be very subtle to predator vision. Nevertheless, according to visual models, larvae should be easily detectable to birds from both herbivore-damaged and control trees. Herbivory affects photosynthetic machinery and light reflectance of leaves, and may thus provide visual foraging cues to birds, although it is not known how these changes appear to birds. We also hypothesized that the changes in leaves may reduce the camouflage of the herbivore. After applying herbivore treatment and using the avian vision models, we found that the leaves of herbivore damage may cause the leaves to appear to birds with higher contrast and greener or a more yellowish colour than control leaves. In addition, although the herbivore was visible to birds, it was slightly less conspicuous when on damaged trees, indicating that the herbivore can be adapted to changes in the food plant. Our results indicate that herbivory causes changes visual properties of leaves, but these changes are likely not obvious to birds.

A novel, 10 kb RNA virus—tentatively named ‘Abisko virus’—was discovered in the transcriptome data of a diseased autumnal moth (Epirrita autumnata) larva, as part of a search for the possible causes of the cyclical nature and mortality associated with geometrid moth dynamics and outbreaks in northern Fennoscandia. Abisko virus has a genome organization similar to that of the insect-infecting negeviruses, but phylogenetic and compositional bias analyses also reveal strong affiliations with plant-infecting viruses, such that both the primary host origin and taxonomic identity of the virus remain in doubt. In an extensive set of larval, pupal, and adult autumnal moth and winter moth (Operophtera brumata) outbreak samples, the virus was only detected in a few adult E. autumnata moths as well as the single larval transcriptome. The Abisko virus is therefore unlikely to be a factor in the Fennoscandia geometrid population dynamics.

The alpine treeline in northern Fennoscandia is composed primarily of mountain birch (Betula pubescens ssp. czerepanovii), a deciduous tree that experiences episodic defoliation due to outbreaks of the autumnal moth (Epirrita autumnata) and winter moth (Operophtera brumata). Here, we use an extensive dendroecological data set to reconstruct historic defoliating outbreaks and relate them to climatic conditions. Our data are from 25 sites in eight valleys in northern Sweden. We used the computer program OUTBREAK to reconstruct moth outbreaks. The reconstructed outbreak record matches the historical record well. There is a significant, but weak relationship between the outbreak severity and temperatures in February, April, July, and August of the outbreak year. Temperatures in the previous May and November were also positively correlated with outbreak severity. For seasonally aggregated temperatures, only autumn temperatures are correlated with outbreak severity. There was no significant correlation...

In addition to climate warming, greater herbivore pressure is anticipated to enhance the emissions of climate-relevant biogenic volatile organic compounds (VOCs) from boreal and subarctic forests and promote the formation of secondary aerosols (SOA) in the atmosphere. We evaluated the effects of Epirrita autumnata, an outbreaking geometrid moth, feeding and larval density on herbivore-induced VOC emissions from mountain birch in laboratory experiments, and assessed the impact of these emissions on SOA formation via ozonolysis in chamber experiments. The results show that herbivore-induced VOC emissions were strongly dependent on larval density. Compared to controls without larval feeding, clear new particle formation by nucleation in the reaction chamber was observed and the SOA mass loadings in the insect-infested samples were significantly higher (up to 150-fold). To our knowledge, this study provides the first controlled documentation of SOA formation from direct VOC emission of deciduous trees damaged by known defoliating herbivores, and suggests that chewing damage on mountain birch foliage could significantly increase reactive VOC emissions that can importantly contribute to SOA formation in subarctic forests. Additional feeding experiments on related silver birch confirmed the SOA results. Thus, herbivory-driven volatiles are likely to play a major role in future biosphere-vegetation feedbacks such as sun-screening under daily 24-hours sunshine in the subarctic.

The microgastrine subset of hymenopteran parasitoids of the geometrid Epirrita autumnata is investigated in Fennoscandia. Ecology, including population dynamics, of the moth has been intensively studied in northern and mountainous Finland, Norway and Sweden. Recently supported hypotheses about the causes of its cyclic population dynamics stress the role of parasitoids, while the parasitoid complex with some 15 species is insufficiently known. The complex includes four solitarymicrogastrine species, Protapanteles anchisiades (Nixon), P. immunis (Wesmael), Cotesia salebrosa (Marshall) and C. autumnatae Shaw, sp. n. Here, we provide detailed figures for the latter, which is morphologically close to C. jucunda (Marshall), and describe the species as new to science. We also providemore general habitus figures of the other three species, as well as an identification key for the four species, aiming to aid recognition of these species by ecologists dealing with microgastrine parasitoids of E. autumnata and their alternative geometrid hosts.

The re-measurement of permanent forest inventories offers a unique opportunity to assess the occurrence and impact of forest disturbances. The present study aims at exploring the main forest damages in Norway based on the extensive data of several consecutive national forest inventories during the period 1995–2014. Five of the most common disturbance agents in Norway are selected for analysis: wind, snow, browsing, fungus and insect damage. The analyses focuses on the frequency and variation along time, the average damage at stand level and the spatial patterns of damage occurrence, resulting in a characterization of the damage produced by disturbances in Norway. The highest damage occurrences by disturbance agent are due to browsing, snow and wind. Snow presents a decreasing temporally trend in damage frequency in the studied period. By forest type, mature and intermediate birch forest are found to be more affected by snow damage, whereas mature spruce forest is by wind damage. The results from this study provide support to the hypothesis that damages by autumnal moth (Epirrita autumnata) on birch are more common in mature stands. No major attacks from bark beetle (Ips typographus) are found, probably related to the lack of major storm damages in the period. Forest types susceptibility to fungus has no apparent variation over time except in the last years, as increased occurrence is observed on mature spruce stands probably correlated with warmer than average periods. Browsing damage causes the most severe losses, as expected, in young stands, and is allocated mainly on the most productive forests. Although some of the disturbances present locally moderate effects, the results show no major disturbances threatening Norwegian forests in the studied period. Finally, the Norwegian national forest inventory demonstrates its reliability as a basis to understand the occurrence and effects of major natural disturbances. We are grateful to the forest doctoral school of the University of Eastern Finland for their kind economic support, as well as to the SIS-Redclim, funded by the Norwegian Institute for Bioeconomy Research.

Summary 1. Many plant species have two inducible indirect defences against herbivores, emission of volatile organic compounds (VOCs) and secretion of extrafloral nectar (EFN). Moreover, herbivore-induced VOCs can serve as airborne signals in plant–plant communication, inducing and/or priming defences in undamaged neighbouring plants. However, it remains largely unknown, particularly in woody plants, whether herbivore-induced VOCs induce and/or prime these two indirect defences simultaneously. 2. We investigated the VOC-mediated induction and priming of VOC emission and EFN secretion in the woody plant hybrid aspen (Populus tremula × Populus tremuloides) in the context of plant–plant communication. Hybrid aspen saplings were exposed to VOCs released either from herbivore-infested or uninfested conspecifics and subsequently challenged by herbivores. VOCs and EFN from exposed plants were collected throughout experiments. Additionally, local and systemic EFN induction was investigated. 3. Exposure to VOCs from infested plants directly induced secretion of EFN sugars but not emission of volatile terpenes. Upon subsequent herbivore attack, however, plants previously exposed to herbivore-induced VOCs released significantly more terpenes than unexposed plants, whereas EFN secretion was not increased, indicating that herbivore-induced VOCs prime VOC emission but not EFN secretion. This switching in the indirect defence priority from induced EFN secretion to primed VOC emission may reflect the potential fitness disadvantages of VOC emission and EFN secretion on their own. In addition, feeding by Epirrita autumnata larvae reduced EFN sugar production both locally and systemically, which disagrees with previous findings and suggests that EFN secretion varies depending on plant and herbivore species. 4. Thereby, our work has parallels and divergences from the most significant previous studies. Nevertheless, it suggests that plants can respond differently to early warning signals and ensuing herbivory and in doing so fine-tune their different indirect defences timely to achieve optimal use of resources.

Plants show rapid adaptation towards resistance against heavy metals. In contrast, tests examining the local adaptation of insect herbivores to heavy metal pollution are lacking. Here we test whether the autumnal moth Epirrita autumnata, living in a heavy metal contaminated area, shows local adaptation in the growth rate and immune function. We gathered male moths from control and polluted areas for use in paternal half-sib crossings in order to ensure that we measure genotypic variation in addition to phenotypic variation. We found genotype × environment interactions in the growth of the larvae suggesting potential local adaptation in the growth of pollution-exposed moths. However, adaptation appears to incur a cost, because we observed reduced performance of the heavy metal adapted strain in a non-polluted environment. Finally, we found that pollution enhanced the immune function in female moths but not in males.

Host plant quality has a major influence on the performance, and ultimately on the fitness of an herbivorous insect, but may also have indirect effects on the third trophic level by affecting an herbivore's defensive ability against natural enemies. In a three-year field study, we examined the effects of natural food quality on the ability of autumnal moths, Epirrita autumnata (Lepidoptera, Geometridae), to defend themselves against parasitoids. In each year, we confirmed the variation in quality of host trees (mountain birch, Betulapubescens ssp. czerepanovii) by determining the mass of pupae reared in mesh bags attached to the trees and the water content of leaves. Individuals grown on high quality trees possessed significantly higher encapsulation rate of a foreign antigen as pupae compared to those on low quality trees during the first and third study years; a parallel trend was also found in the second study year, although this difference was not statistically significant. However, in spite of observed differences in encapsulation rates, individuals reared on high and low quality trees did not differ in their levels of parasitisation when exposed to hymenopteran parasioids in the wild and thus were equally vulnerable. Accordingly, the encapsulation response seems not to play a major role on the population ecology scale in the studied system. Our findings also stress the importance of direct resistance tests, which should be conducted along with tests of insect immune function.

1. The abundance and fecundity-related body size variation of the cyclic autumnal moth Epirrita autumnata were monitored from the early increase phase and throughout the outbreak to the end of the density decline in northernmost Norway during 1999-2006. Another geometrid, the winter moth Operophtera brumata, did not increase in density until the autumnal moth had its post-peak in 2004, and was at its own peak concurrent with the steeply declining autumnal moth abundance in 2005-06. 2. The body size variables measured (forewing lengths of males and females and hind femur lengths of males) of the autumnal moth showed a similar density-dependent response, i.e. increasing density was associated with decreasing body size and fecundity. Nevertheless, regression analyses clearly ranked the pooled geometrid abundance without a time lag as the best predictor for the body size variation, ahead of the abundance of the autumnal moth or past abundance of all geometrids. 3. Nondelayed effects of lowered food quality and absolute shortage of foliage under congested conditions are the most plausible reasons for reduced body size. 4. Two most commonly proposed causal factors of the autumnal moth population cycle, i.e. delayed inducible resistance of the host plant (mountain birch Betula pubescens czerepanovii) and delayed density-dependent parasitism by specialized hymenopteran parasitoids, cannot easily explain the diverging population trends between the autumnal and winter moths. 5. We suggest that either the inducible resistance of the host tree or the host utilization of the most important parasitoids and/or pathogens have to be strictly species-specific between these closely related moth species to produce the population dynamics observed. That fecundity of the autumnal moth was best related to the pooled geometrid abundance weakens support for the former hypothesis, while our lack of host-specific information limits conclusions about the role of natural enemies.

Previously, we found that Epirrita autumnata larvae, which were fed with leaves that came from a metal polluted area, had an enhanced immune function. To investigate further the cause of the enhanced immunity we examined experimentally the direct influence of the heavy metals, copper and nickel, on the immune function of the geometrid moth we artificially added the metals to the surface of birch leaves to examine their direct effects. As a measurement of the strength of immune function, we used encapsulation rate against a nylon monofilament. A moderate amount of Ni and Cu in the diet of moth larvae increased their encapsulation rate, but a large amount of Cu caused the immune function to decrease. The results indicate that Ni and Cu have direct effects on immunity. Moderate amounts of those metals increase immune function, but large amounts lead to inhibition of immune function.

Plants use volatile organic compounds to attract invertebrate predators and parasitoids of their herbivore pests. Recently, it has been suggested that plants, either through visual or olfactory cues, may also "cry for help" from vertebrate predators such as birds. We show that in a laboratory choice test, passerine birds (Parus major and Cyanistes caeruleus) were attracted to the intact branches of trees (Betula pendula) suffering from foliar damage caused by herbivore larvae (Epirrita autumnata) in nontest branches. Species, age, or sex of the experimental bird or lighting (ultraviolet [UV] or non-UV) did not affect the preference. However, the birds made a clear choice between the treatments when the trees came from a forest patch receiving more sunlight, whereas no obvious choice was observed when the trees came from a shadier forest patch. Results of the choice test were supported by the spectral reflectance of tree leaves. In the sunnier forest patch, control trees reflected more visible light than the herbivore trees, whereas no such difference was found in the shadier forest patch trees. We suggest that avian predators use their vision within visible wavelengths to find insect-rich plants even when they do not see the prey items or damaged leaves. Copyright 2008, Oxford University Press.

Fluctuating asymmetry (FA) has been suggested as a useful indicator of elevation stress and, hence, distribution limits in plants. However, no plant studies have been carried out to test (i) whether FA shows a gradual increase towards the alpine distribution limit and (ii) whether FA responds to elevation stress independent of other stressors which is necessary for FA to be a useful indicator in this context. To test these two hypotheses, this 2-year field study investigated the dose–response relationship between elevation stress and FA in mountain birch (Betula pubescens) under contrasting levels of insect attack in northern Norway. The results showed that FA increased linearly from sea level towards the tree line in both years independent of insect attack, which had no observable effect on FA, i.e. insect attack did not appear to disturb the FA-elevation relation. Thus, in mountain birch, FA appeared to be a reliable indicator of elevation stress. Further investigation is now needed in order to develop this hypothesis.

1. In recent studies, it has been argued that sunspot activity forces the Epirrita autumnata 9-10-year outbreak periodicity in the mountain birch forest of Fennoscandia. For the following reasons, we challenge this conclusion. 2. With a 10-year outbreak cycle of E. autumnata and the 11-year sunspot cycle, it is expected that the cycles will run in-phase, out-of-phase and in-phase within 10 x 11 years. Hence, given such cycle lengths, sunspot activity should not affect outbreak periods. For a test, the E. autumnata series should be at least 110 years in length. 3. A well-documented E. autumnata outbreak series of 81 years (1888-1968; outbreak periods IV-XII) exists. This series is here lengthened to 114 years by adding outbreak frequencies for three decades (1969-2001). 4. By lengthening the series, three more E. autumnata/Operophtera brumata periods (XIII, XIV, XV) are identified. Period XV, like several earlier periods, was of the moving type, i.e. outbreaks moved in a wavelike manner from northern Fennoscandia to southern Norway. 5. As with several earlier outbreak periods in central northern Fennoscandia, the main timing of periods XIII-XV centred at the middle of the decades. In contrast, outbreaks at the extreme north-western coast of Norway centred at the decadal shifts, i.e. about 1979, 1989 and 1999. Supported by historical documents, we explain the 1979 and 1999 outbreaks as the final expressions of east-west outbreak waves that branched off from the main waves which moved southward during periods XIII and XV. These side-waves in the north are new observations. Outbreaks at the decadal shift 1989/1990 may have been of a more complex nature. 6. We find that sunspot activity does not explain outbreak waves. Furthermore, a test of our 114-year long E. autumnata series against the contemporaneous sunspot series shows that the two series run in-phase and out-of-phase. The observed interval between the two cycles coming in-phase agrees with the expected interval. This challenges the hypothesis of sunspot synchronization of the E. autumnata (and O. brumata) outbreaks.

The potential capacity of larval growth and immune response traits of the autumnal moth to adapt to heavy metal polluted environment was tested experimentally. Both the relative growth rate (RGR) and pupal weight were significantly higher in control trees than on polluted trees, indicating that metal pollution prevented the insect from achieving maximal growth on birch leaves. Larval growth rates of different broods differed significantly between metal contaminated and control birches. However, pupal weight of broods, which is considered more important for fitness than growth rate, in response to pollution did not differ. Immune response was significantly higher in moths exposed to pollution than in moths that were exposed to control environment suggesting that pollution enhances the immune defense of defoliators. Encapsulation rate tended to differ between broods indicating that the immune function has potential to respond to selection.

The autumnal moth (Epirrita autumnata) is a cyclically outbreaking forest Lepidoptera with circumpolar distribution and substantial impact on Northern ecosystems. We have isolated 21 microsatellites from the species to facilitate population genetic studies of population cycles, outbreaks, and crashes. First, PCR primers and PCR conditions were developed to amplify 19 trinucleotide loci and two tetranucleotide loci in six multiplex PCR approaches and then analyzed for species specificity, sensitivity and precision. Twelve of the loci showed simple tandem repeat array structures while nine loci showed imperfect repeat structures, and repeat numbers varied in our material between six and 15. The application in population genetics for all the 21 microsatellites were further validated in 48 autumnal moths sampled from Northern Norway, and allelic variation was detected in 19 loci. The detected numbers of alleles per locus ranged from two to 13, and the observed and expected heterozygosities varied from 0.04 to 0.69 and 0.04 to 0.79, respectively. Evidence for linkage disequilibrium was found for six loci as well as indication of one null allele. We find that these novel microsatellites and their multiplex-PCR assays are suitable for further research on fine- and large-scale population-genetic studies of Epirrita autumnata. tri- and tetranucleotide microsatellites; multiplex PCR; Lepidoptera; population genetics

Outbreaks by geometrid moths periodically cause mass mortality of trees and state changes in understorey vegetation in sub-arctic mountain birch forest in northern Scandinavia. In order to assess the short-term impacts of such disturbance on forest bird communities, we took bird censuses in forest where almost all birch trees had been killed by moth outbreaks 2–4 years before the study and in undamaged forest. The study was repeated in two locations (Kirkenes and Tana) with contrasting forest structure and fragmentation. Using a hierarchical community model, we show that the total abundance of birds in Kirkenes was only about 25 % lower in damaged than undamaged forest and that species richness differed even less between the two forest types. Meanwhile, neither bird abundance nor species richness differed between damaged and undamaged forest in Tana. The observed patterns in abundance were mainly driven by a few very common species. Only a single species showed indication of being more abundant in damaged forest. Thus, our findings indicate that bird communities in sub-arctic mountain birch forest have a high degree of resistance to forest damage caused by moth outbreaks. We suggest that bird populations in outbreak-affected forest may be maintained by surviving trees and by standing dead tree trunks, which help maintain the vertical structure of the forest habitat. The fact that many of the studied bird species are habitat generalists may also explain their apparently weak responses to the damage caused by the outbreak. Our results do not point towards forest damage caused by moth outbreaks as a major driver of change in bird communities in the study system, although more long-term research is needed to substantiate this conclusion.

Many species of forest lepidopterans exhibit regular population cycles, which culminate in outbreak densities at approximately ten-year intervals. Population peaks and mass outbreaks typically occur synchronously and may lead to extensive forest damages over large geographic areas. Here, we report patterns of spatial synchrony among cyclic autumnal moth (Epirrita autumnata) populations across Fennoscandia, as inferred from 24 long-term (10–33 years) data sets. The study provides the first formal analysis of spatial synchrony of this pest species which damages mountain birch (Betula pubescens ssp. czerepanovii) forests in the sub Arctic. We detected positive cross-correlations in population growth rates between the time series, indicating overall spatial synchrony. However, we found the strongest degree of synchrony within geographically and climatically distinct regional clusters, into which time series were partitioned using cluster analyses. Within regional clusters, moth populations were exposed to the synchronizing effects of common, spatially autocorrelated environmental conditions, i.e. a Moran effect. Consequently, we conclude that a geographically and climatically restricted Moran effect, perhaps interacting with dispersal, is the most likely explanation for the regionally partitioned pattern of synchrony among autumnal moth populations in Fennoscandia. Our results emphasize that high amounts of environmental variation may result in a clear structuring of spatial synchrony at unexpectedly small scales.

During population outbreaks, top-down and bottom-up factors are unable to control defoliator numbers. To our knowledge, details of biotic interactions leading to increased population density have not been studied during real population outbreaks. We experimentally assessed the strength of plant defenses and of insect immunocompetence, assumed to contribute to active insect resistance against parasitoids and pathogens, in the geometrid Epirrita autumnata during a steep increase in population density. We demonstrated rapid (same-season) induced resistance in the foliage of its host, mountain birch. The response was systemic, spreading throughout the tree, and retarded larval growth rate by approximately 10%. On the other hand, no direct delayed carry-over effects were found in the next season in larval growth rate, mortality, or pupal mass. Larval damage to a tree during the previous year, however, significantly (by approximately 13%) accelerated the advance of the immune response (measured as melanization of an implant inserted into the pupal hemocoel). The encapsulation rate correlated positively with larval mortality in trees in which larvae had been introduced the previous year, but not in control trees. Both of these observations suggest that induced plant defense was associated with an increased insect immunocompetence during the population increase.

• Aims The aim of the study was to examine oxidative responses in subarctic mountain birch, Betula pubescens subsp. czerepanovii, induced by herbivory and manual wounding. • Methods Herbivory-induced changes in polyphenoloxidase, peroxidase and catalase activities in birch leaves were determined. A cytochemical dye, 3,3-diaminobenzidine, was used for the in situ and in vivo detection of H2O2 accumulation as a response to herbivory and wounding. To localize peroxidase activity in leaves, 10 mm H2O2 was applied to the dye reagent. • Key Results Feeding by autumnal moth, Epirrita autumnata, larvae caused an induction in polyphenoloxidase and peroxidase activities within 24 h, and a concomitant decrease in the activity of antioxidative catalases in wounded leaves. Wounding also induced H2O2 accumulation, which may have both direct and indirect defensive properties against herbivores. Wound sites and guard cells showed a high level of peroxidase activity, which may efficiently restrict invasion by micro-organisms. • Conclusion Birch oxidases together with their substrates may form an important front line in defence against herbivores and pathogens.

The two forest-defoliating geometrid moth species Operophtera brumata and Epirrita autumnata are known to exhibit different altitudinal distribution patterns in northern birch forests. One possible explanation for this is that altitudinal climatic variation differentially affects the performance of two species through mismatching larval and host plant phenology. We explored this hypothesis by investigating the relationship between larval phenology and leaf phenology of Betula pubescens, which is the main host plant of both moth species, along ten replicate altitudinal transects during two springs with contrasting climate in northern Norway. There was a distinct monotonous cline in host plant phenology with increasing altitude in both years of the study, but the development of the leaves were generally 14 days later in the first of the 2 years due to cold spring weather. We found that larval development of both species closely tracked host plant leaf phenology independent of altitude and year. However, at the time of sampling, E. autumnata was approximately one instar ahead of O. brumata at all altitudes, probably reflecting that E. autumnata has faster early instar growth than O. brumata. The abundance of O. brumata was lowest at the altitudinal forest-line, while E. autumnata was lowest near sea level. Our results do not indicate that the altitudinal distribution patterns of the two moth species is due to any phenological mismatch between larval and host plant phenology. We suggest rather that natural enemies at low altitudes limit larval survival and thus abundance of E. autumnata, while an early onset of winter at the forest limit reduces survival of late eclosing adults of O. brumata.

Short-lived insect herbivores should be able to adapt to the resistance mechanisms of their long-lived woody hosts because the life span of a single host will encompass numerous generations of herbivores. However, adaptation may be slowed down if host genotypes can create, in a single genotype, such large phenotypic variation in traits relevant for the herbivore that it matches variance among host genotypes. We tested this hypothesis by measuring leaf consumption by, and growth of, half-sibs of the geometrid moth Epirrita autumnata on individual birch trees, during three instars. The instar � /tree interaction, rather than tree identity alone, was a significant variance component for both consumption and growth, indicating that different larval instars ranked individual trees differently. Both consumption and growth varied most between the 3rd and the later (4th and 5th) instars, coinciding with rapid seasonal changes in numerous nutritive and phenolic traits of maturing leaves. Thus, developmental variance in the leaf quality of individual trees may reduce the likelihood of E. autumnata genotypes adapting to the defenses of their host trees. We did not find evidence of in the ability of different half-sibs to utilize individual trees or leaf stages, indicating that E. autumnata larvae are generalists over a wide variety of host traits.

Analysis of the mitochondrial DNA (mtDNA) control region (CR) was used to examine the dispersal of females of a geo- metrid moth, Epirrita autumnata, in Fennoscandia. A 542-bp-portion of the CR of 200 individuals from four northern and four southern localities was sequenced. The mtDNA CR of E. autumnata contains a substantial amount of variation as a total of 108 mtDNA haplotypes were observed. Between the northern and the southern localities (~1100 km), there was a moderate level of genetic differentiation (FST = 0.128). The amount of variation in the mtDNA CR of E. autumnata was lower in the north than in the south. The reduction in genetic variability may result from a combination of historical bottlenecks that date back to the post-glacial recolonization of Fennoscandia and, present-day bottlenecks due to the northern E. autumnata populations experiencing repeated outbreaks followed by collapse in population size. On a small spatial scale (0.6-19 km), within the northern and southern areas, no genetic differentiation was detected suggesting ongoing gene flow due to the dispersal of E. autumnata females among the localities. This finding was contrary to our earlier expectation of poor flying ability of E. autumnata females.

It is often claimed that some organisms maximise individual growth rates (within limits set by physiological and environmental constraints) whereas others do not. Adequate experimental protocol to test for these hypotheses has not been available. We proceeded from the idea that a maximiser should be unable to further improve its growth performance, i.e. there should be no adaptive plasticity in relative growth rates. In the case of an insect larva always maximising its weight gain, we should thus expect no increase in growth rate following a short-term environmental perturbation. Another prediction is that initial and final weights of a larval instar should be highly correlated. We applied this approach to test for the growth rate maximiser status of the geometrid moth Epirrita autumnata. Based on various lines of ecological evidence it had previously been suggested that this spring-feeding species is strongly selected for high growth rates, and may be growing at its maximum physiological potential. Contrary to expectations, the larvae responded to starvation treatments by a subsequent compensatory increase in their relative growth rates. Such a plasticity may indicate that the rates of gaining weight are not maximised even in a ‘classical’ example a time-constrained organism. This observation supports the idea that endogenous regulation of growth rates is widespread, and it remains to be shown if true maximisation ever exists in nature. In contrast, possibilities to compensate for adverse conditions experienced early in larval development appeared to be limited. This was indicated by the strong correlation between initial weight in the last instar, and pupal weight. Though consistent with our expectations, such an inability to compensate can hardly be ascribed to growth rate maximisation, as indicated by quantitatively similar patterns found in an unrelated species with different ecology, the butterfly Polygonia c-album.

Variation in in situ growth performance of the mountain birch as indicated by the widths of annual rings was analysed and related mainly to temperature and herbivory using ring width series from five heath forest sites in the Lake Tornetrask area, northern Sweden. Climate explained 48-64% of the variation in age-corrected mean ring width series. In general, the effect of current year July followed by June temperature was most important at all sites. A warm May resulted in wider rings due to an earlier budburst. Short-term (inter-annual) responses to increased temperature were in most cases not reflected into long-term responses (decades). A large proportion of the variation in stem mean ring width was due to variation among stems within trees (81%) in these polycormic trees, while variation among sites was marginal (0.4%). Within trees, main stems grew faster and were more responsive to climate variation than subordinate stems. No effect of insect herbivory on ring width was found at low defoliation levels ( ≤ 12%). At a defoliation level of ca 84% a one-year reduction in stem growth was observed while the growth reduction (ca 50% reduction in ring width) lasted for 4 yr after ca 93% defoliation. After outbreaks resulting in complete defoliation and some stem mortality, ring widths of surviving stems mainly responded with increased growth. Basal sprouts, emerging just after a severe insect outbreak with a high mortality of old stems, grew faster than sprouts occurring during other periods. It is concluded that the mountain birch is well adapted to recover from Epirrita outbreaks; the ability to produce basal sprouts, that can benefit from an existing root system for fast initial growth, is one important mechanism for this. P S. Karlsson (staffan.karlsson@ebc.uu.se), Dept of Plant Ecology, EBC, Uppsala Univ, Villavagen 14, SE-752 36 Uppsala, Sweden. - O. Tenow and H. Bylund, Dept of Entomology, Swedish Univ. of Agricultural Sciences, P.O. Box 7044, SE-750 07 Uppsala, Sweden. - J. Hoogesteger, Kuninkaantie 856, FIN-47450 Perheniemi, Finland - M. Weih, Dept of Short Rotation Forestry, Swedish Univ. of Agricultural Sciences, P.O. Box 7016, SE-750 07 Uppsala. Sweden.

In northern Fennoscandia, the geometrid moths Epirrita autumnata and Operophtera brumata have cyclicities in density with mass occurrence at 10-year intervals. The larvae of Epirrita and Operophtera attain a size of 2‐3 cm and 1.5‐2 cm, respectively, and are nutritious food items for passerine birds. To examine whether these larvae have any numerical and/or functional influence on a passerine bird community (mountain birch forest in Budal, central Norway) during a 30-year period (1972‐2001), I estimated their abundance (number of larvae per 100 sweeps) in the birch canopy, and the densities of breeding birds in the passerine community. In addition, from 1972 to 1998, I monitored the nesting success of five of the bird species. The foraging pattern of the most abundant bird species and their gizzard contents (adults and nestlings) were examined in 1972‐78 (covering population peaks of both the geometrids). Population peaks of Epirrita occurred in 1975‐76, 1985‐86 and 1996, and of Operophtera in 1976‐77, 1986‐87 and 1997‐98. The passerine community consisted of eight species that were territorial in all 30 years, one species in 26 years, three species in 14‐21 years and three species in 1‐4 years. Only the Brambling Fringilla montifringilla population responded numerically to the fluctuations of Epirrita and Operophtera. Brambling was also the only species whose mean clutch size varied between years, and this correlated positively with the density of Epirrita. The mean annual nesting success of Willow Warbler Phylloscopus trochilus, Bluethroat Luscinia svecica and Common Redpoll Carduelis flammea tended to be higher in years with mass outbreaks of Epirrita, but was significantly so only for Reed Bunting Emberiza schoeniclus. The abundance of Operophtera larvae showed no influence on the nesting success of any bird species. The passerines foraged more frequently in the birch canopy in the Epirrita outbreak years (1975‐76) than in the years before or after. Gizzard analyses of five adult passerine species and their nestlings showed that Epirrita was the main food item in 1974‐76. Even though Operophtera occurred in large numbers in birch trees in 1976 and 1977, only a few larvae were found in the gizzards of the passerines. None of the passerines showed an increase in their population density in the year following the larval outbreaks, but the densities of Willow Warbler and Bluethroat increased in the succeeding year, indicating a higher return rate for these species. The study shows the existence of a dietary response and also indicates a reproductive response to the changes in the abundance of Epirrita in mountain birch forest. The lack of numerical response in the passerines (except the Brambling) to the fluctuation in Epirrita contrasts with the pattern described for passerine communities in northern temperate deciduous forests in North America, where Lepidoptera caterpillars periodically have mass outbreaks.

The breeding passerine bird community of 9 km2 of a primeval subalpine birch forest in Swedish Lapland was censused yearly from 1963 through 1999. The community density remained stable, fluctuating around a mean value of 403 ± 85 territories/km2. Species richness increased by an average of one species every four years. Twenty species made up 98% of the community and Phylloscopus trochilus (40%) and Fringilla montifringilla (19%) remained the dominant species. The species turnover rate between successive seasons was on average 30%. The mean temperatures in June and July were positively correlated to the community density two years and one year later, respectively, less so for the densities of the individual species. Fringilla montifringilla increased in density during the first two years of each Epirrita autumnata caterpillar outbreak, whereas the other species responded only weakly or not at all. All species tended to fluctuate in parallel (no significant opposite trends) which together with the other results suggests that the investigated bird community is far from saturated.

In Lepidoptera, effects of larval crowding on life history traits may manifest themselves as changes in growth rate and duration of larval period. For the outbreaking geometrid moth, Epirrita autumnata, impacts of crowding have been shown in earlier laboratory studies, and these responses were modified by diet quality. In this study, the aim was to find out the specific nature of crowding effects of E. autumnata in the field. E. autumnata larvae were reared individually and in groups in mesh bags on mountain birch, both on good and poor quality foliage. Three field experiments were carried out with different densities in the crowded treatment (12, 21 and 45 larvae per 100 short shoots of mountain birch), simulating intermediate to severe outbreaks. The study revealed the density interval (12-21) within which impacts of crowding become evident. Significant effects were mostly found with the two highest densities in the crowded treatment combined with poor foliage quality. In those treatments, crowding resulted in 8-12% and 24-33% decrease in pupal mass and larval survival, respectively. The results of the present field experiments, however, did not corroborate the results of earlier laboratory studies: duration of larval period was not substantially affected, net effects of crowding were negative and interactions between crowding and foliage quality were contrary to those obtained in earlier laboratory studies. In many other Lepidoptera as well, the knowledge on responses to crowding may be mostly qualitative since most crowding experiments have been conducted in laboratory.

Shading is assumed to reduce allocation to plant phenolics and to defense in general. We here report the results of experimental shading of individual branches or whole canopies in mountain birch on foliar chemistry and on the growth and consumption of a geometrid, Epirrita autumnata. Branch-wide shading tended to have at least as strong effects on both leaf chemistry and herbivore performance as canopy-wide shading, indicating local responses of the host to shading. Responses to shading varied among the key leaf traits. Leaf water content was higher and toughness lower in shaded than in non-shaded leaves. Leaf sugars were lower and protein-bound and free amino acids higher in shaded than in control leaves. Sucrose and galactose were at high levels in unshaded branches adjacent to shaded ones, suggesting that partial shading enhanced translocation of sugars within canopies. Total phenolics and soluble proanthocyanidins were low in both shading treatments. Of the other phenolic groups, concentrations of gallotannins and cell-wall-bound proanthocyanidins did not differ between shaded and non-shaded leaves. Epirrita larvae grew better in both types of shading treatments compared to either unshaded control trees or to unshaded branches in the branch-shading trees. By far the most important correlate of larval growth was the amount of water consumed with leaf mass (r = 0.94). When variance in water intake was standardized (also largely eliminating parallel variation in proteins), fructose and glucose still had significant positive correlations and proanthocyanidins negative with larval growth on control but not on shade leaves. Concentrations of several phenolic compounds correlated negatively with intake of dry matter and especially water, and different phenolics were important in shaded (gallotannins) and in control (flavonoids) leaves. Our findings strongly suggest that the effects of putatively defensive leaf traits on insect consumption and growth interact with nutritive leaf traits, particularly with water.

Habitats are now becoming increasingly fragmented throughout the world due to intense cultivation. As a consequence, populations of some animals with low mobility have become isolated, thus increasing the risk of inbreeding and local extinction. In Britain, weakly flying geometric moths of the genus Epirrita are a good model species with which to test the genetic effects of habitat fragmentation on insect populations. Genetic variation within and between populations of two Epirrita species captured using a network of light traps at two spatial scales (local and national) was assessed using allozyme electrophoresis, with particular reference to the local scale (the 330-ha arable farm estate at Rothamsted, Hertfordshire, in southern Britain). Populations sampled widely in England and Wales displayed low (but statistically significant) levels of genetic differentiation for both species (F st = 0.0051–0.0114 and 0.0226 for E. dilutata and E. christyi , respectively). However, analysis of large samples of E. dilutata from four small woods at Rothamsted revealed low (F st = 0.0046) but significant differentiation, indicating that gene flow was restricted, even at this very small scale. It was concluded that small intervening patches of farmland (often a few fields width) were enough to prevent genetic homogeneity. The close similarity between more distant Epirrita populations was considered to be a result of historical, rather than recurrent gene flow, as genetic equilibrium between drift and gene flow is unlikely over such scales. © 2003 The Linnean Society of London. Biological Journal of the Linnean Society , 2003, 78, 467–477.

The mast depression hypothesis has been put forward to explain the 9- to 10-year population cycle of the autumnal moth (Epirrita autumnata; Lepidoptera: Geometridae) in northern Fennoscandia. We analysed long-term data from Finnish Lapland in order to evaluate the critical assumption of the mast depression hypothesis: that better individual performance of herbivores, followed by high annual growth rate of populations, occurs in the year following mast seeding of the host, the mountain birch ( Betula pubescens ssp. czerepanovii). Since mast seeding has been suggested to occur at the expense of chemical defence against herbivores, we bioassayed the quality of birch leaves from the same trees by means of yearly growth trials with autumnal moth larvae. We also measured the size of wild adults as a determinant of potential fecundity of the species in different years. The relative growth rate of larvae was poorer in post-mast years compared to other years, rather than better as assumed by the hypothesis. Conversely, a slight indication of the increase in potential fecundity was observed due to the somewhat larger adult size in post-mast years. Population growth rate estimates, however, showed that the increase in fecundity would have to be much higher to facilitate population increase towards a cyclic peak with outbreak density. Accordingly, our two data sets do not support the assumption of a higher annual growth rate in autumnal moth populations subsequent to mast seeding of the host, thereby contradicting the predictions of the mast depression hypothesis. Temperatures, when indexed by the North Atlantic Oscillation and accumulated thermal sums, were observed to correlate with the abundance or rate of population change of the autumnal moth. The factors underlying the regular population cycles of the autumnal moth, however, remain unidentified. Overall, we suggest that the causal agents in cyclic insect population dynamics should be clarified by field experimentation, since trophic interactions are complex and are further modified by abiotic factors such as climate.

Several studies have found genetic variation in plant resistance to herbivory. One of the explanations suggested for the observed intermediate levels of resistance are the costs of resistance, i.e., negative genetic correlations between resistance and other fitness components that may constrain the evolution of resistance. We studied the cost of herbivore resistance by investigating the genetic correlations between resistance traits and plant growth traits, and between resistance to insect and mammalian herbivores in cloned saplings of silver birch, Betula pendula. We used the performance of a geometrid moth, Epirrita autumnata, as an indicator of insect resistance. The numbers of resin droplets at the base and at the tip of the saplings correlate with mammalian resistance, and were thus used here as indicators of vole and hare resistance, respectively. We have previously observed genetic variation in these resistance traits. Further, we examined the correlations between several groups of secondary chemicals and plant growth traits. Finally, to reveal the effect of environmental factors on the trade-offs mentioned above, we investigated the correlations in saplings that were grown at two nutrient levels. We found significant negative correlations between indices of constitutive insect resistance and relative height growth in non-fertilized saplings, indicating cost of constitutive insect resistance. The two groups of secondary chemicals that have been shown to correlate strongly with constitutive insect resistance, i.e., condensed tannins and flavonol glycosides (especially myricetin glycosides), had different genetic correlations with plant traits; the concentration of condensed tannins did not correlate negatively with any of the plant traits, whereas the concentration of flavonol glycosides correlated negatively with plant height. Insect and mammalian resistance did not correlate negatively, indicating no ecological trade-offs.

The organization of the mitochondrial DNA (mtDNA) control region (CR) of the autumnal moth, Epirrita autumnata, is described. The E. autumnata CR presents a distinct type of lepidopteran CR with domains of non-repetitive and repetitive sequences. The CRs show considerable length variation owing to a variable number of short ~29-bp sequence blocks that are repeated between 6 and 14 times in tandem. The organization of such a tandem array is unique among the insect CRs examined so far. Furthermore, the E. autumnata CR, which may reach 1075 bp in length, is considerably longer than previously reported lepidopteran CRs, which reach 311–499 bp in length. Like other lepidopteran CRs, the E. autumnata CR contains two long homopolymer runs that may be involved in mtDNA replication and (or) transcription.Key words: Lepidoptera, Epirrita autumnata, mitochondrial DNA, control region, tandem array.