Waves and synchrony in Epirrita autumnata/Operophtera brumata outbreaks. I. Lagged synchrony: regionally, locally and among species

1. In 1990-2003, during a complete 10-year outbreak cycle, the synchrony of the birch defoliating outbreaks of the geometrids Epirrita autumnata and Operophtera brumata was studied quantitatively in the northern part of the Fennoscandian mountain chain (the Scandes). Data were supplemented with similar data from 1964 to 1966 and historical information. A 30-year series of field data from one locality in southern Scandes made possible interregional comparisons. 2. In 1991, outbreaks started in north-eastern Fennoscandia and moved westward like a wave and reached the outer coast of north-western Norway in about 2000. This wave is a new observation. In the same years, a previously documented outbreak wave moved southward along the Scandes. 3. Outbreak periods have usually occurred around the middle of each decade. Seemingly unrelated population peaks at the decadal shift 2000 were reported from islands at the coast of north-western Norway. They are shown here to have been the final ripples of the east-west wave. 4. At some localities, O. brumata peaked 2 years after E. autumnata. A lag of 1 or 2 years also occurred at the locality in southern Scandes. This interspecific time lag is a new observation. In accordance with the north-south wave, a time-lag of 1-2 years occurred between the fluctuations of northern and southern E. autumnata and O. brumata populations. 5. The population peak of E. autumnata occurred 1 year earlier at one locality than at a nearby locality. This pattern and particular altitudinal shifts of the O. brumata population density at these localities repeated in two outbreak periods. This indicates that, for example, local climate may modify outbreak synchrony between nearby localities. 6. At the same localities, O. brumata peaked first at one altitude and 1 or 2 years later at another altitude. This vertical lag is a new observation. 7. E. autumnata shows fluctuation traits similar to some other cyclic animals, e.g. the larch budmoth in the European Alps, some European tetraonid birds and the Canadian snow-shoe hare. These similarities (and dissimilarities) in intra- and interspecific synchronies and causes of E. autumnata and O. brumata synchronies, regionally, locally and among the two species are discussed.     

Waves and synchrony in Epirrita autumnata/Operophtera brumata outbreaks. II. Sunspot activity cannot explain cyclic outbreaks

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.     

Shifting altitudinal distribution of outbreak zones of winter moth Operophtera brumata in sub‐arctic birch forest: a response to recent climate warming?

Climatic change is expected to affect the extent and severity of geometrid moth outbreaks, a major disturbance factor in sub-arctic birch forests. Previous studies have reported that the two geometrid species involved, autumnal moth and winter moth, differ in their temperature requirements and, consequently, in their altitudinal and latitudinal distribution patterns. In this study, we document the altitudinal distribution of winter moth outbreaks in a large coastal area in northern Norway. We show that, in the present winter moth outbreak, defoliated birch stands were seen as distinct zones with a rather constant width in the uppermost part of the forest and where the upper limit coincided with the forest line. The outbreak zone closely followed the spatially variable forest line as an undulating belt, although some of the variation in outbreak zone width was also related to variation in topographical variables, such as distance from the coast, forest line altitude, and slope of the terrain. A distinct outbreak zone at the altitudinal forest line is the typical picture that has been depicted in more qualitative historical records on previous outbreaks of autumnal moth rather than winter moth. We suggest that the recent documented climate warming in this region may have induced a shift in distribution of the winter moth both relative to topography and geography. Further investigation is, however, required to substantiate these suspicions.     

Anisotropic patterned population synchrony in climatic gradients indicates nonlinear climatic forcing

Although climatic forcing has been suspected to be the most common cause of spatial population synchrony owing to the Moran effect, it has proved difficult to disentangle the impact of climate from other possible causes of synchrony based on population survey data. Nonlinear population responses to climatic variation may be a part of this difficulty, but they can also provide an opportunity to highlight the climate impacts through targeted survey designs. In particular, when species distribution ranges encompass consistent spatial gradients in climate (e.g. according to latitude or altitude), such gradients can be strategically included in the spatial design of population surveys as to facilitate comparisons of spatial synchrony patterns across and along the gradient. In that case, we predict that nonlinear impacts of climatic variation on population growth rates will result in anisotropic (direction specific) synchrony patterns in the sense that synchrony will drop faster with distance along the climatic gradient than across it. We provide an empirical case study to exemplify survey design and analyses. Of two sympatric species of geometrids, inhabiting an altitudinal gradient in subarctic birch forest, one (Operophtera brumata L.) showed anisotropic synchrony consistent with a strongly nonlinear sensitivity to climatic variation, whereas the other (Epirrita autumnata Bkh.) did not. These results are interpreted in light of the biological characteristics of the species.     

Geographically partitioned spatial synchrony among cyclic moth populations

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.     

Rapid northwards expansion of a forest insect pest attributed to spring phenology matching with sub‐Arctic birch

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 (LDT_A.a.=4.71 °C, LDT_O.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.     

Ecosystem Impacts of a Range Expanding Forest Defoliator at the Forest-Tundra Ecotone

Insect outbreaks in northern-boreal forests are expected to intensify owing to climate warming, but our understanding of direct and cascading impacts of insect outbreaks on forest ecosystem functioning is deficient. The duration and severity of outbreaks by geometrid moths in northern Fennoscandian mountain birch forests have been shown to be accentuated by a recent climate-mediated range expansion, in particular of winter moth (Operophtera brumata). Here, we assess the effect of moth outbreak severity, quantified from satellite-based defoliation maps, on the state of understory vegetation and the abundance of key vertebrate herbivores in mountain birch forest in northern Norway. We show that the most recent moth outbreak caused a regional-scale state change to the understory vegetation, mainly due to a shift in dominance from the allelopathic and unpalatable dwarf-shrub Empetrum nigrum to the productive and palatable grass Avenella flexuosa. Both these central understory plant species responded significantly and nonlinearly to increasing outbreak severity. We further provide evidence that the effects of the outbreak on understory vegetation cascaded to cause strong but opposite impacts on the abundance of the two most common herbivore groups. Rodents increased with defoliation, largely mirroring the increase in A. flexuosa, whereas ungulate abundance instead showed a decreasing trend. Our analyses also suggest that the response of understory vegetation to defoliation may depend on the initial state of the forest, with poorer forest types potentially allowing stronger responses to defoliation.     

Outbreaks by canopy-feeding geometrid moth cause state-dependent shifts in understorey plant communities

The increased spread of insect outbreaks is among the most severe impacts of climate warming predicted for northern boreal forest ecosystems. Compound disturbances by insect herbivores can cause sharp transitions between vegetation states with implications for ecosystem productivity and climate feedbacks. By analysing vegetation plots prior to and immediately after a severe and widespread outbreak by geometrid moths in the birch forest-tundra ecotone, we document a shift in forest understorey community composition in response to the moth outbreak. Prior to the moth outbreak, the plots divided into two oligotrophic and one eutrophic plant community. The moth outbreak caused a vegetation state shift in the two oligotrophic communities, but only minor changes in the eutrophic community. In the spatially most widespread communities, oligotrophic dwarf shrub birch forest, dominance by the allelopathic dwarf shrub Empetrum nigrum ssp. hermaphroditum, was effectively broken and replaced by a community dominated by the graminoid Avenella flexuosa, in a manner qualitatively similar to the effect of wild fires in E. nigrum communities in coniferous boreal forest further south. As dominance by E. nigrum is associated with retrogressive succession the observed vegetation state shift has widespread implications for ecosystem productivity on a regional scale. Our findings reveal that the impact of moth outbreaks on the northern boreal birch forest system is highly initial-state dependent, and that the widespread oligotrophic communities have a low resistance to such disturbances. This provides a case for the notion that climate impacts on arctic and northern boreal vegetation may take place most abruptly when conveyed by changed dynamics of irruptive herbivores.     

Phenological diversity in the interactions between winter moth (Operophtera brumata) larvae and parasitoid wasps in sub-arctic mountain birch forest

Population cycles of the winter moth (Operophtera brumata) in sub-arctic coastal birch forests show high spatiotemporal variation in amplitude. Peak larval densities range from levels causing little foliage damage to outbreaks causing spatially extensive defoliation. Moreover, outbreaks typically occur at or near the altitudinal treeline. It has been hypothesized that spatiotemporal variation in O. brumata cycle amplitude results from climate-induced variation in the degree of phenological matching between trophic levels, possibly between moth larvae and parasitoids. The likelihood of mismatching phenologies between larvae and parasitoids is expected to depend on how specialized parasitoids are, both as individual species and as a guild, to attacking specific larval developmental stages (i.e. instars). To investigate the larval instar-specificity of parasitoids, we studied the timing of parasitoid attacks relative to larval phenology. We employed an observational study design, with sequential sampling over the larval period, along an altitudinal gradient harbouring a pronounced treeline outbreak of O. brumata. Within the larval parasitoid guild, containing seven species groups, the timing of attack by different groups followed a successional sequence throughout the moth's larval period and each group attacked 1-2 instars. Such phenological diversity within parasitoid guilds may lower the likelihood of climate-induced trophic mismatches between victim populations and many/all of their enemies. Parasitism rates declined with increasing altitude for most parasitoid groups and for the parasitoid guild as a whole. However, the observed spatiotemporal parasitism patterns provided no clear evidence for or against altitudinal mismatch between larval and parasitoid phenology.     

Effects of host-plant shift on immune and other key life-history traits of an eruptive Geometrid, Epirrita autumnata (Borkhausen)

Abstract.  1. Population density of Epirrita autumnata (Lepidoptera: Geometridae) reaches outbreak densities regularly in northernmost Scandinavia. During these outbreak years, the most abundant host species, the mountain birch ( Betula pubescens ssp. czerepanovii ), is regularly exhausted, although larvae may rescue themselves from starvation by using alternative host species.
2. In this paper, the effects of the shift of host species on the immune defence and other life-history traits of E. autumnata were investigated, and possible consequences for population dynamics were briefly discussed. Moth larvae were reared on the leaves of the main host, mountain birch, until larvae reached their third instar. After this, larvae were allocated randomly to five treatments: larvae were either allowed to finish larval stage on the mountain birch or were shifted onto four alternative host species that are typical species for the area.
3. As expected, the host species had a major effect on fitness traits: body weight, development, and survival rate of the moths. The pupal weight was lower and development rates slower on the three alternative host species, Salix myrsinifolia Salisb., Vaccinium uliginosum L., and Betula nana L., than on the main host, mountain birch.
4. The immunity was, however, the same or better on the alternative hosts than on the main host. The immunity and pupal weights were negatively related, suggesting a trade-off between body size and immunocompetence.
5. The decreased body size and fecundity of E. autumnata during outbreak years may be partly due to the shift to alternative host species whereas the host-plant species probably does not affect markedly the rate of parasitism.     

Spatial responses of two herbivore groups to a geometrid larva on mountain birch

Direct or plant-mediated interactions between herbivores may modify their spatial distribution among and within plants. In this study, we examined the effect of a leaf-chewing geometrid, the autumnal moth (Epirrita autumnata), on two different herbivore groups, leaf rolling Deporaus betulae weevils and Eriocrania spp. leafminers, both feeding on mountain birch (Betula pubescens ssp. czerepanovii). The exact locations of herbivores within tree canopies were mapped during three successive summers. In the first 2 years, some trees were artificially colonized by eggs of the autumnal moth to induce both rapid and delayed resistance in the foliage. The natural infection levels of the pathogenic rust fungus (Melampsoridium betulinum), potentially involved in species interactions, were also recorded. At the level of the whole tree, the density of D. betulae leaf rolls was lower in trees infested by the autumnal moth in the same year. However, the feeding locations within trees were partly segregated: D. betulae favoured shadier branches, while E. autumnata preferred the sunny parts of the canopy. The autumnal moth did not affect current- or following-year density of leafminers at the tree or branch level. Trees infected by rust had fewer leafminers in the same summer than noninfected trees. There were no interaction effects between defoliation by the autumnal moth and rust infection, and no delayed effects on the abundance of other herbivores the following year. Taken together, these findings suggest that the autumnal moth has a negative, partially plant-mediated impact on D. betulae, and can reduce the extent of current-year defoliation caused by D. betulae. This may be beneficial for the mountain birch, since the greater part of D. betulae damage occurs around or after the end of the larval period of the autumnal moth, which may be a critical time for tree recovery after moth outbreaks.     

Fecundity of the autumnal moth depends on pooled geometrid abundance without a time lag: implications for cyclic population dynamics

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.     

Wound-induced oxidative responses in mountain birch leaves

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.     

Stand age-structure influence in a low population peak of Epirrita autumnata in a mountain birch forest

Population densities of Epirrita autumnata caterpillars were recorded from 1984 to 1990 m four mountain birch stands of two age categories, young and old Caterpillar populations peaked m 1986–87, and peak densities were higher in the old than in the young stands Recorded peak densities were low compared with synchronous populations with outbreak densities in neighbouring areas Differences in caterpillar densities between young and old stands could not be explained by differences in survival of early instar caterpillars or in parasitoid-caused larval mortality Instead, tree-age related differences in amounts of suitable oviposition sites and possible changes in food quality due to ageing and lower vigour of mature trees are suggested to explain why E autumnata populations tend to be higher in old stands     

Specialist and generalist natural enemies as an explanation for geographical gradients in population cycles of northern herbivores

Within Fennoscandia, two well-studied groups of herbivores exhibit clear geographical gradients in their population dynamics. Populations of a forest lepidopteran ( Epirrita autumnata , the autumnal moth) and voles of the genera Microtus and Clethrionomys show pronounced multi-annual cycles in the north but become more stable towards the south. Here we review empirical and theoretical studies on these species, mainly regarding the biological mechanisms that are assumed to generate the pattern of population dynamics in both systems. We conclude that the specialist/generalist predation hypothesis offers a common explanation for the population cycles and their geographical gradients irrespective of whether a herbivorous insect or small mammals are concerned. According to this hypothesis, originally developed for the Fennoscandian voles, but now applied also to E. autumnata , population cycles are generated by specialist natural enemies (predators for the voles and parasitoids for E. autumnata ). Furthermore, the dynamic shift from cycles to stability is assumed to be caused by an increase in the density and diversity of generalist natural enemies from north to south in Fennoscandia.     

Crowding-induced responses in a geometrid moth revisited: a field experiment

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.     

Phenology and abundance in relation to climatic variation in a sub-arctic insect herbivore–mountain birch system

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.     

Invading and resident defoliators in a changing climate: cold tolerance and predictions concerning extreme winter cold as a range‐limiting factor

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.     

Metabolic modifications of birch leaf phenolics by an herbivorous insect: detoxification of flavonoid aglycones via glycosylation

The metabolic modifications of birch (Betula pubescens Ehrh.) leaf phenolics in the digestive tract of its major defoliator, larvae of the autumnal moth Epirrita autumnata, were studied. The main phenolic acids of birch, i.e. chlorogenic and p-coumaroylquinic acids, were isomerised in the alkaline digestive tract. Moreover, only 16 to 92% of the ingested amounts of chlorogenic acid were found in the faeces of individual larvae; the missing portion is possibly being used in the formation of reactive o-quinones. Water-soluble flavonoid glycosides were mostly excreted unaltered. In contrast, lipophilic flavonoid aglycones were not excreted as such, but as glycosides after being detoxified by E. autumnata via glycosylation. When the larvae were fed with leaf-painted acacetin and kaempferide, i.e. two naturally occurring birch leaf flavonoid aglycones, acacetin-7-O-glucoside and kaempferide-3-O-glucoside appeared in larval faeces as major metabolites. However, the efficiency of aglycone glycosylation varied-, ranging from 17 to 33%, depending on the aglycone and its dietary level. There was also large variation in the efficiency of glycosylation--from 2 to 57%--among individual larvae. These results demonstrate a compound-specific metabolism of phenolic compounds, leading to different phenolic profiles in the insect gut compared to its leaf diet.