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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.

Red autumn colouration of trees is the result of newly synthesized anthocyanin pigments in senescing autumn leaves. As anthocyanin accumulation is costly and the trait is not present in all species, anthocyanins must have an adaptive significance in autumn leaves. According to the coevolution hypothesis of autumn colours, red autumn leaves warn herbivorous insects – especially aphids that migrate to reproduce in trees in the autumn – that the tree will not be a suitable host for their offspring in spring due to a high level of chemical defence or lack of nutrients. The signalling allows trees to avoid herbivores and herbivores to choose better host trees. In this study the coevolution hypothesis was tested with four deciduous tree species that have red autumn leaf colouration – European aspen (Populus tremula L.) (Salicaceae), rowan (Sorbus aucuparia L.) (Rosaceae), mountain birch [Betula pubescens ssp. czerepanovii (NI Orlova) Hamet‐Ahti], and dwarf birch (Betula nana L.) (Betulaceae), and with two generalist herbivores, the autumnal moth [Epirrita autumnata (Borkhausen)] and the winter moth [Operophtera brumata (L.)] (both Lepidoptera: Geometridae). Anthocyanin concentrations of autumn leaves were determined from leaf samples and the growth performance parameters of the moth larvae on the study trees were measured in the spring. Trees with higher anthocyanin concentration in the autumn were predicted to be low‐quality food for the herbivores. Our results clearly showed that anthocyanin concentration was not correlated with the growth performance of the moths in any of the studied tree species. Thus, our study does not support the coevolution hypothesis of autumn colours.

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.

Large-scale spatial synchrony is common in population dynamics. In nearly all empirical data sets, population synchrony decays with distance. Large-scale spatial population synchrony of 20 common moth species was studied using 367 separate light-trap time series of 3–27 years, at 13–19 localities in southern Finland. The maximum distance between trap sites was ca 1075 km. The degree of spatial synchrony in three weather variables (daily mean and minimum temperature, and daily total precipitation) from 15 weather stations was also studied. Data were analysed with spatial non-parametric correlation functions (Sncf). Statistically significant large-scale spatial synchrony in population fluctuations occurred in 85% of the species (Scotopteryx chenopodiata, Eulithis populata, Chloroclysta citrata, Epirrita autumnata, Perizoma alchemillata, P. didymata, Eupithecia pusillata, Macaria wauaria, Itame brunneata, Cabera pusaria, C. exanthemata, Eilema lurideola, Rusina ferruginea, Amphipoea fucosa, Orthosia gothica, Cerapteryx graminis and Diarsia mendica), as well as in all weather variables. Spatial synchrony decreased with distance (=distance decay) in 45% of the species, and there was no indication of distance decay in the remaining species. Climate change is likely to cause an increasing frequency of extreme weather events, and insects are strongly affected by such environmental stochasticity. In combination with the high frequency of large-scale spatial synchrony, which did not show distance decay as a rule in these data, even currently common moth species may become threatened in the near future.

Some insectivorous birds orient towards insect-defoliated trees even when they do not see the foliar damage or the herbivores. There are, however, only a few studies that have examined the mechanisms behind this foraging behaviour. Previous studies suggest that birds can use olfactory foraging cues (e.g. volatile organic compounds (VOCs) emitted by defoliated plants), indirect visual cues or a combination of the two sensory cues. VOCs from insect-defoliated plants are known to attract natural enemies of herbivores, and researchers have hypothesized that VOCs could also act as olfactory foraging cues for birds. We conducted three experiments across a range of spatial scales to test this hypothesis. In each experiment, birds were presented with olfactory cues and their behavioural responses or foraging outcomes were observed. In the first experiment, two different VOC blends, designed to simulate the volatile emissions of mountain birch (Betula pubescens ssp. czerepanovii) after defoliation by autumnal moth (Epirrita autumnata) larvae, were used in behavioural experiments in aviaries with pied flycatchers (Ficedula hypoleuca). The second experiment was a field-based trial of bird foraging efficiency; the same VOC blends were applied to mountain birches, silver birches (B. pendula) and European white birches (B. pubescens) with plasticine larvae attached to the trees to serve as artificial prey for birds and provide a means to monitor predation rate. In the third experiment, the attractiveness of silver birch saplings defoliated by autumnal moth larvae versus intact controls was tested with great tits (Parus major) and blue tits (Cyanistes caeruleus) in an aviary. Birds did not orient towards either artificial or real trees with VOC supplements or towards herbivore-damaged saplings when these saplings and undamaged alternatives were hidden from view. These findings do not support the hypothesis that olfactory foraging cues are necessary in the attraction of birds to herbivore-damaged trees.

‘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.

In birch, Betula pubescens, herbivore-induced delayed induced resistance (DIR) of defoliated trees may cause a strong reduction in the potential fecundity of a geometrid folivore Epirrita autumnata. In this study, we examined the biochemical basis of DIR in birch leaves during a natural outbreak of E. autumnata. A set of experimental trees was defoliated at four sites by wild larvae in the peak year of the outbreak, whereas control trees were protected from defoliation by spraying with an insecticide. The biochemical composition of leaves was analysed in the following year and, although the DIR response was weak during this outbreak, causing less than a 20% reduction in the potential fecundity of E. autumnata, some consistent relationships between defoliation, biochemistry and pupal mass of E. autumnata suggested a general biochemical basis for the defoliation-induced responses in birch leaves. Total concentrations of nitrogen, sugars and acetone-insoluble residue (e.g. cell wall polysaccharides, cell-wall-bound phenolics, protein, starch, lignin and hemicellulose) were consistently lower, and total concentrations of phenolics, especially of gallotannins and soluble proanthocyanidins, were higher in the leaves of trees defoliated in the previous year than in those protected from defoliation. The capacity of tannins to precipitate proteins correlated with contents of gallotannins, and was highest in defoliated trees. The pupal mass of E. autumnata showed a strong, positive correlation with concentrations of nitrogen and sugars, and a negative correlation with the acetone-insoluble residue and gallotannins in foliage. Correlations with other measured biochemical traits were weak. The correlation coefficients between biochemical traits and pupal mass consistently had similar signs for both defoliated and insecticide-sprayed trees, suggesting that variation in leaf quality due to defoliation in the previous year was based on similar biochemical traits as variation for other reasons. We suggest that DIR is associated with reduced growth activity of leaves, and may be seen as a delay in the biochemical maturation of leaves in defoliated trees. This explains the high concentration of gallotannins in defoliated trees, a characteristic feature of young leaves. However, the lower content of nitrogen and the higher content of soluble proanthocyanidins in defoliated trees are traits usually characterising mature, not young, leaves, indicating defoliation-induced changes in chemistry in addition to modified leaf age. Our results emphasise the importance of understanding the natural changes in chemistry during leaf maturation when interpreting defoliation-induced changes in leaf biochemistry.

Due to rapidly changing physical and biochemical characteristics of growing leaves, correlations between traits of foliage biochemistry and the performance indices of flush feeding herbivores may vary considerably following relatively minor changes in experimental conditions. We examined the effects of the seasonal and inter-tree variation of a comprehensive array of biochemical compounds on the success of an early season geometrid, Epirrita autumnata, feeding on maturing foliage of mountain birch, Betula pubescens ssp. czerepanovii. We monitored the concentrations of individual phenolics, sugars, total nitrogen, nitrogen of proteins, and nitrogen of soluble compounds, water and acetone-insoluble residue. Simultaneously we recorded larval consumption, physiological performance, growth, and pupal mass of E. autumnata. We found significant phenological changes in almost all leaf traits measured. In bioassays with half-grown leaves, leaf gallotannin concentrations showed a nonlinear effect: in trees with high foliar gallotannin concentrations (over 10 mg g−1), physiological performance was strongly reduced by high gallotannin concentrations. In trees with lower gallotannin concentrations, on the other hand, larval growth was reduced by soluble proanthocyanidins, not gallotannins. Differences between high and low gallotannin trees largely depended on phenology, i.e., on the age of leaves. However, not all the differences in leaf traits between late (with high gallotannin concentrations at the time of the bioassay) and early flushing trees disappeared with leaf maturation, indicating that there is also phenology-independent variance in the tree population. In the full-grown leaves of all the study trees, low concentrations of water and of nitrogen of proteins (but not nitrogen of soluble compounds) were the main factors reducing pupal masses of E. autumnata, while neither gallotannin nor proanthocyanidins now played a significant role. The observed change in the factors underlying leaf quality (from gallotannins and proanthocyanidins to nitrogen and water) relate to the activity of the shikimate pathway and the formation of cell walls: gallotannins and proanthocyanidins are both produced in the pathway, and these tannins are assumed to contribute – via binding into cell walls – to tough and durable cell walls. Interestingly, low quality of leaves did not automatically translate into low foliar consumption (i.e., benefits to the tree). On the trees with young, high gallotannin leaves, larvae actually increased consumption on low quality foliage. In the group of trees with slightly more developed, low gallotannin leaves, the quality of leaves did not clearly modify amounts consumed. In full-grown leaves, low leaf quality strongly reduced leaf consumption. These results emphasize the strong influence of tree phenology on the relationships between biochemical compounds and the herbivore.

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...

The principal objective of the study was to characterize effects of increasing density of the autumnal moth, Epirrita autumnata (Lepidoptera, Geometridae), for 3 successive years on the change in metabolome of mountain birch trees (Betula pubescens ssp. czerepanovii). During the three study years (2000–2002), the larval density varied from low to very high (outbreak) density. Samples of leaves were collected from the same 12 trees at the same phenological stage of the trees each year. The leaves were collected from undamaged trees in the first year, from trees slightly damaged by larvae in the second year, and from trees heavily damaged by larvae in the third year. Metabolome analysis showed that the increase in density of E. autumnata larvae and degree of damage of birch trees caused multiple biochemical changes in the leaves, including increased concentrations of phenolic compounds (proanthocyanidins and hydrolysable tannins) and reduced concentrations of nutritive metabolites (monosaccharides, amino acids and some organic acids). These changes reduced the quality of leaves as food for larvae and, probably, were associated with induced chemical resistance of the birches to herbivorous insects. Additionally, the concentration of α-tocopherol was significantly higher in both slightly and heavily damaged trees. Mechanisms of changes in the metabolism of phenolic compounds and carbohydrates, and the role of α-tocopherol in their regulation are discussed.

In 2001, Hamilton and Brown proposed a controversial hypothesis of handicap signalling to potential insect parasites as an adaptive explanation for autumn leaf colouration. In subsequent studies there has been little attention to the costs and benefits of early autumnal colour change. Yet, in an observational study by Hagen et al. (2003) it was demonstrated that birch trees [Betula pubescens ssp. czerepanovii Ehrhart (Betulaceae)] turning yellow early in autumn had less damage from insects chewing on leaves the subsequent summer. Here, two experiments are presented which test the mechanisms in this model. The first addresses the proposed defence of leaves of B. pubescens ssp. czerepanovii by letting caterpillars of Epirrita autumnata Borkhausen (Lepidoptera: Geometridae), the birches’ most common insect parasites, choose between leaves from trees that either turned yellow late or early the foregoing autumn. The second experiment addresses whether adult female E. autumnata choose between early or late senescent (i.e., yellow or green) ‘twigs’ when ovipositing in autumn. We could not find evidence of preferences in either larvae or females, suggesting that timing of colour change in B. pubescens ssp. czerepanovii is not a warning signal that elicits a response in E. autumnata.

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

Earlier studies have suggested that insectivorous birds, similar to invertebrate predators and parasitoids, may be guided by herbivore-induced plant volatiles (HIPVs) to damaged, herbivore-rich trees. Recent studies have also shown that birds use olfaction more than previously thought, underlying the potential for HIPVs to be sensed by insectivorous birds and utilised during foraging for prey. The HIPV production in plants is mediated, at least partly, by the jasmonic acid signalling pathway, and similar HIPVs to those induced by herbivores can often be induced by exposing plants to methyl jasmonate (MeJa). We studied the effects of MeJa on volatile emission and bird attraction using mature mountain birches (Betula pubescens ssp. czerepanovii) under natural conditions in northern Finland. Experimental trees were assigned to four treatment groups: herbivore-damaged [autumnal moth (Epirrita autumnata)], higher dose of MeJa (30 mM), lower dose of MeJa (15 mM) and control. All trees had three branches covered with mesh bags, but there were larvae inside the bags only of the herbivore-damage treatment. Bird predation rate was monitored with artificial plasticine larvae which were checked daily for peck marks. Birds most often pecked the larvae in the herbivore-damaged trees, but the attractiveness of MeJa-treated trees did not differ from the control. High within-treatment variation in systemic HIPV emissions probably masked MeJa treatment effects. The bird predation rate was high in birches that emitted large amounts of α-pinene. Thus, α-pinene may be one cue used by birds to find herbivore-rich birches.

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.

1 The induced resistance of the subarctic mountain birch Betula pubescens ssp. czerepanovii is a well-characterized phenomenon, whereas the induced responses of Betula nana L., one of the parental species of mountain birch, have not yet been characterized. Betula nana is more resistant to several classes of insectivorous herbivores than the mountain birch, although the mechanisms responsible for the better ability to resist herbivores are not known. 2 The present study aimed to determine the metabolic changes that are induced by early season herbivory in B. nana leaves and to study the effects of rapidly induced resistance on the growth of Epirrita autumnata larvae. 3 Defoliation of B. nana was accomplished by E. autumnata larvae and leaf samples for chemical analyses were collected when the defoliating larvae were at their third and fifth instar. At the same time, laboratory assays for the growth and consumption rates of E. autumnata larvae were conducted. 4 The wounding of leaves by E. autumna larvae induced the production of ellagitannins (ETs) in B. nana. Intriguingly, the concentrations of protein-bound amino acids were also induced by herbivory; however, an increase in proteins was not mirrored in the growth rate of larvae, which was less on the induced foliage. The decreased growth rate of larvae was apparently linked to the increased concentrations of oxidatively-active ETs and the high concentration of ETs may explain the better resistance of this parental species compared with the hybrid mountain birch with its lower levels of ETs.

1. Studies of insect communities rarely support the parasitoid–host regulation hypothesis. Spatio-temporal variation in parasitoid prevalence due to complex food web interactions or abiotic factors may prevent parasitoids from regulating hosts. 2. We examined the relative contribution of spatial (altitude) and temporal (years) sources to total variation in parasitoid prevalence rates in outbreaks of Epirrita autumnata Borkhausen and Operophtera brumata Linnaeus populations. We tested whether prevalence rates of generalist parasitoids were correlated between sympatric host populations and to what extent any of the parasitoids were host density dependent. 3. Four larval parasitoids (two specialists and two generalists) exhibited significantly structured spatio-temporal dynamics over years and altitudes. The prevalence rates of one of the generalists were spatio-temporally correlated between the two host species, while for the other they were not. 4. Three parasitoids showed tendencies for direct or delayed positive density dependence as expected from numerical and functional responses to their hosts. However, the effects were weak and minute compared to the variation attributed to year and altitude. 5. We conclude that unknown aspects of the larval parasitoid ecology that co-vary with altitude and year in the study system dominate their prevalence dynamics and thus act to hinder density-dependent responses that could potentially regulate host populations.

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.

Another theoretical and practical issue related to autumn leaf coloration is the issue of leaf color variability. While several authors proposed that such variability in timing and intensity of autumn leaf coloration has been selected for (Archetti 2000; Hamilton and Brown 2001; Hagen et al. 2003; Ougham et al. 2005; Sinkkonen 2006a; Sinkkonen et al. 2012), theoretically, this variability may in certain cases be a non-defensive adaptation, or not adaptive at all. Variation in the timing of color change may reflect positional constraints, such as the position of leaves within the tree, the level of exposure to sunlight or wind action, the status of the tree as dominant, sub-dominant or suppressed individual within the population, or being an understory or canopy species, or else reflect physiological pressures such as fluctuations in water supply or mineral nutrition, factors that may significantly influence a tree’s performance irrespective of its genotype (Kozlowski et al. 1991; Kozlowski and Pallardy 1997). The results of Hagen et al. (2003) that in Betula pubescens ssp. czerepanovii trees that turn yellow early in autumn (September) was negatively correlated with insect damage the following season, did not match the results of Isaksen and Folstad (2014) for the same subspecies but in different years. Isaksen and Folstad (2014) found no difference in the attacks of the most common caterpillar species Epirrita autumnata on leaves, or in the oviposition by mature females on twigs senessing early or late. Thus, it seems that at least for that taxon, the difference in succeptability to herbivory is not only related to variability in phenology, but is also year and habitat depended. Isaksen and Folstad (2014) rejected the aposematic function in that case. However, at least for various Finnish Betula pendula (silver birch) genotypes, Sinkkonen et al. (2012) showed that autumn leaf coloration varies significantly among genotypes, and that this influences fitness and evolution (Lev-Yadun and Keasar 2012). In that species, genotypes that expressed the strongest leaf reflectance in early autumn harbored more egg-laying females of the specialist aphid Euceraphis betulae than those expressing strong leaf reflectance several weeks later. Silfver et al. (2015) studied the phenology of shoot growth termination in various Finnish Betula pendula genotypes that differ in that character. Genotypes that terminated shoot growth earlier suffered from higher aphid egg loads after a short, but not after a long autumn. The Sinkkonen et al. (2012) and Silfver et al. (2015) studies illuminated the need to document the variability in autumn leaf coloration and the corresponding aphid attacks in many other taxa and under various environmental conditions.

The exposure of plants to abiotic stresses like air pollutants may increase their susceptibility to herbivores. However, abiotic stresses often induce the accumulation of phenolic compounds that may have adverse effects on plant-eating animals. In this study, we examined the effects of long-term (14 years) deposition of simulated acid rain (H2SO4) and heavy metals (CuNi) on the plant–herbivore interaction. The used species were well-known model species of plant–herbivore interaction, namely the mountain birch (Betula pubescens ssp. czerepanovii) and autumnal moth (Epirrita autumnata). The study set-up was factorial with four combinations of two treatments: −acid−CuNi, +acid−CuNi, −acid+CuNi, and +acid+CuNi. The deposition of pollutants had no marked effects on the growth, survival, or immune function of the autumnal moth although the chemistry of birch leaves was markedly affected. The concentrations of gallic acid, chlorogenic acid, three of hydrolyzable tannins (HTs), and quercetin glycosides were induced by the acid rain treatment when compared to leaves not treated with acid rain. We concluded that an increase in compounds with high redox capacity was a defense against oxidative stress caused by pollution treatments. We suggested that pollution might have increased the deposition of non-phenolic antioxidants like ascorbate that diminish the adverse effects of phenolics on herbivores. We also stated that abiotic stresses might play an equally strong role in the evolution of phenolics than herbivory if not larger.

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.

1. Winter temperatures in northern latitudes are predicted to increase markedly as a result of ongoing climate change, thus making the invasion of new insect defoliators possible. The establishment of new outbreak pest species may have major effects on northern ecosystems that are particularly sensitive to disturbances. 2. Effects of winter minimum temperatures under field and laboratory conditions were examined and limitations by minimum temperatures on future range expansion were investigated for invasive [Operophtera brumata (Lepidoptera: Geometridae)] and potentially invasive [Agriopis aurantiaria (Lepidoptera: Geometridae)] birch-feeding forest pests. The results for the studied invasive and potentially invasive moths were compared with the parameters of the resident moth species Epirrita autumnata (Lepidoptera: Geometridae). 3. The results showed tolerated critical temperatures of the invader (O. brumata) and the resident (E. autumnata) were more similar (differing only by 1 °C), whereas the potential invader (A. aurantiaria) was much less tolerant of cold temperatures. Although describing different stages of overwintering, results were consistent between laboratory and field studies except for those at one field location, at which other abiotic conditions are suggested to have significant influence on moth egg survival. 4. Based on the present results and expected changes in winter temperatures over the next 30 years, the range expansion of an established invasive species may be predicted. No limitations were found regarding the possible future invasion of a new pest species to northern Fennoscandia. The importance of studying a species' whole overwintering period is highlighted and further studies devoted to the effects of other abiotic factors in addition to the effects of temperature are suggested.

Indirect plant defence mechanisms enhance the effectiveness of natural enemies of herbivores. Herbivore-induced plant volatiles (HIPVs) attract the parasitoids of insect herbivores as shown both in numerous choice tests conducted under laboratory conditions and in relatively few common-garden setups in agro-ecosystems. However, the importance of this indirect defence trait at higher levels of biological organization has yet to be investigated through natural field experiments. Here, we report a field experiment of larval parasitism of two cyclic geometrid defoliators in herbivore-damaged and fairly intact mountain birch Betula pubescens ssp. czerepanovii under natural conditions. Parasitism rates in larvae of the autumnal (Epirrita autumnata) and winter moth (Operophtera brumata) exposed for 30 h on defoliated trees were more than twice as high as those on control trees. This finding indicates that hymenopteran parasitoids were attracted to previously defoliated trees by some cues from the host plants, HIPVs being the most likely candidates. The third trophic level should thus be considered in natural plant herbivore interactions. Furthermore, parasitoids and food resources are key factors in the population regulation of forest insect pests, and indirect plant defences could be important in their interactions. Our research also emphasizes the quality of control treatments in field experiments, since immediate plant responses easily obscure the results as soon as control trees become infested by herbivorous insects.

Species range displacements owing to shifts in temporal associations between trophic levels are expected consequences of climate warming. Climate-induced range expansions have been shown for two irruptive forest defoliators, the geometrids Operophtera brumata and Epirrita autumnata, causing more extensive forest damage in sub-Arctic Fennoscandia. Here, we document a rapid northwards expansion of a novel irruptive geometrid, Agriopis aurantiaria, into the same region, with the aim of providing insights into mechanisms underlying the recent geometrid range expansions and subsequent forest damage. Based on regional scale data on occurrences and a quantitative monitoring of population densities along the invasion front, we show that, since the first records of larval specimens in the region in 1997–1998, the species has spread northwards to approximately 70°N, and caused severe defoliation locally during 2004–2006. Through targeted studies of larval phenology of A. aurantiaria and O. brumata, as well as spring phenology of birch, along meso-scale climatic gradients, we show that A. aurantiaria displays a similar dynamics and development as O. brumata, albeit with a consistent phenological lag of 0.75–1 instar. Experiments of the temperature requirements for egg hatching and for budburst in birch showed that this phenological lag is caused by delayed egg hatching in A. aurantiaria relative to O. brumata. A. aurantiaria had a higher development threshold (LDTA.a.=4.71 °C, LDTO.b.=1.41 °C), and hatched later and in less synchrony with budburst than O. brumata at the lower end of the studied temperature range. We can conclude that recent warmer springs have provided phenological match between A. aurantiaria and sub-Arctic birch which may intensify the cumulative impact of geometrid outbreaks on this forest ecosystem. Higher spring temperatures will increase spring phenological synchrony between A. aurantiaria and its host, which suggests that a further expansion of the outbreak range of A. aurantiaria can be expected.

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.

We used fine-spatial resolution remotely sensed data combined with tree-ring parameters in order to assess and reconstruct disturbances in mountain birch ( Betula pubescens ) forests caused by Epirrita autumnata (autumnal moth). Research was conducted in the area of Lake Tornetrask in northern Sweden where we utilized five proxy parameters to detect insect outbreak events over the 19th and 20th centuries. Digital change detection was applied on three pairs of multi-temporal NDVI images from Landsat TM/ETM+ to detect significant reductions in the photosynthetic activity of forested areas during disturbed growing seasons. An image segmentation gap-fill procedure was developed in order to compensate missing scan lines in Landsat ETM+ “SLC-off” images. To account for a potential dependence of local outbreak levels on elevation, a digital elevation model was included in the defoliation recognition process. The resulting damage distribution map allowed for the assessment of outbreak intensity and distribution at the stand level and was combined with tree-ring data and historical documents to produce a multi-evidence outbreak detection. Defoliation events in the tree-ring data were recognized as significant deviations from temperature related growth. Our outbreak detection scheme allowed for the reconstruction of nine major insect outbreaks over the past two centuries. The reconstruction proved reliable but only robust for severe defoliation events. Low-intensity incidents were not captured.

The individual and combined effects of elevated CO 2 and 0 3 on the foliar chemistry of silver birch (Betula pendula Roth) and on the performance of five potential birch-defoliating insect herbivore species (two geometrid moths, one lymantrid moth and two weevils) were examined. Elevated CO 2 decreased the water concentration in both short- and long-shoot leaves, but the effect of CO 2 on the concentration of nitrogen and individual phenolic compounds was mediated by 0 3 treatment, tree genotype and leaf type. Elevated 0 3 increased the total carbon concentration only in short-shoot leaves. Bioassays showed that elevated CO 2 increased the food consumption rate of juvenile Epirrita autumnata and Rheumaptera hastata larvae fed with short-and long-shoot leaves in spring and mid-summer, respectively, but had no effect on the growth of larvae. The contribution of leaf quality variables to the observed CO 2 effects indicate that insect compensatory consumption may be related to leaf age. Elevated CO 2 increased the food preference of only two tested species: Phyllobius argentatus (CO 2 alone) and R. hastata (CO 2 combined with 0 3 ). The observed stimulus was dependent on tree genotype and the measured leaf quality variables explained only a portion of the stimulus. Elevated 0 3 decreased the growth of flush-feeding young E. autumnata larvae, irrespective of CO 2 concentration, apparently via reductions in general food quality. Therefore, the increasing tropospheric 0 3 concentration could pose a health risk for juvenile early-season birch folivores in future. In conclusion, the effects of elevated 0 3 were found to be detrimental to the performance of early-season insect herbivores in birch whereas elevated CO 2 had only minor effects on insect performance despite changes in food quality related foliar chemistry.

1 One of the unresolved questions in studies on population dynamics of forest Lepidoptera is why some populations at times reach outbreak densities, whereas others never do. Resolving this question is especially challenging if populations of the same species in different areas or of closely-related species in the same area are considered. 2 The present study focused on three closely-related geometrid moth species, autumnal Epirrita autumnata, winter Operophtera brumata and northern winter moths Operophtera fagata, in southern Finland. There, winter and northern winter moth populations can reach outbreak densities, whereas autumnal moth densities stay relatively low. 3 We tested the hypothesis that a lower vulnerability to pupal predation may explain the observed differences in population dynamics. The results obtained do not support this hypothesis because pupal predation probabilities were not significantly different between the two genera within or without the Operophtera outbreak area or in years with or without a current Operophtera outbreak. 4 Overall, pupal predation was even higher in winter and northern winter moths than in autumnal moths. Differences in larval predation and parasitism, as well as in the reproductive capacities of the species, might be other candidates.

When plants are damaged by herbivorous insects they emit a blend of volatile organic compounds (VOCs) which include a range or terpenoids and green leaf volatiles (GLVs) formed via different metabolic pathways. The precise timing of these emissions upon the onset of herbivore feeding has not been fully elucidated, and the information that is available has been mainly obtained through laboratory based studies. We investigated emissions of VOCs from Populus tremula L. xP. tremuloides Michx. during the first 20 h of feeding by Epirrita autumnata (autumnal moth) larvae in a field site. The study was conducted using Proton Transfer Reaction-Mass Spectrometry (PTR-MS) to measure emissions online, with samples collected for subsequent analysis by complementary gas chromatography-mass spectrometry for purposes of compound identification. GLV emission peaks occurred sporadically from the outset, indicating herbivore activity, while terpene emissions were induced within 16 h. We present data detailing the patterns of monoterpene (MT), GLV and sesquiterpene (SQT) emissions during the early stages of herbivore feeding showing diurnal MT and SQT emission that is correlated more with temperature than light. Peculiarities in the timing of SQT emissions prompted us to conduct a thorough characterization of the equipment used to collect VOCs and thus corroborate the accuracy of results. A laboratory based analysis of the throughput of known GLV, MT and SQT standards at different temperatures was made with PTR-MS. Enclosure temperatures of 12, 20 and 25 degrees C had little influence on the response time for dynamic measurements of a GLV or MT. However, there was a clear effect on SQT measurements. Elucidation of emission patterns in real-time is dependent upon the dynamics of cuvettes at different temperatures.

Immature leaves of birches and other deciduous trees and shrubs are the principal food resource for the larvae of the autumnal moth, Epirrita autumnata (Borkhausen) (Lepidoptera: Geometridae). However, the larvae of this geometrid, which displays pronounced population cycles and causes massive forest defoliations in northernmost Europe, are also frequently found on the catkins of mountain birch, Betula pubescens ssp. czerepanovii (Orlova) Hamet-Ahti (Betulaceae). To examine whether autumnal moth larvae visit birch catkins for their high nutritional value, we conducted two growth experiments in which the larval diet was supplemented with male catkins, and a single test where female catkins were used. We found that the presence of male catkins in the larval diet had a positive effect on the larvae, in relation to their pupal mass (reflecting life span fecundity), survival probability, as well as the duration of the larval period. The presence of female catkins in the diet did not affect larval performance. Our findings suggest that young autumnal moth larvae actively move to male catkins, which provide a nutritional benefit and thus lead to a better performance compared with those feeding elsewhere. Another possible benefit for young larvae choosing the catkins instead of leaves may involve predator or parasitoid avoidance.