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 701N, and caused severe defoliation locally during [2004][2005][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. 5 4.71 1C, LDT O.b. 5 1.41 1C), 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.

K. 2003. Effects of host shading on consumption and growth of the geometrid Epirrita autumnata: interactive roles of water, primary and secondary compounds. -Oikos 103: 3 -16.Shading is assumed to reduce allocation to plant phenolics and to defense in general. We here report the results of experimental shading of individual branches or whole canopies in mountain birch on foliar chemistry and on the growth and consumption of a geometrid, Epirrita autumnata. Branch-wide shading tended to have at least as strong effects on both leaf chemistry and herbivore performance as canopy-wide shading, indicating local responses of the host to shading. Responses to shading varied among the key leaf traits. Leaf water content was higher and toughness lower in shaded than in non-shaded leaves. Leaf sugars were lower and protein-bound and free amino acids higher in shaded than in control leaves. Sucrose and galactose were at high levels in unshaded branches adjacent to shaded ones, suggesting that partial shading enhanced translocation of sugars within canopies. Total phenolics and soluble proanthocyanidins were low in both shading treatments. Of the other phenolic groups, concentrations of gallotannins and cell-wall-bound proanthocyanidins did not differ between shaded and non-shaded leaves. Epirrita larvae grew better in both types of shading treatments compared to either unshaded control trees or to unshaded branches in the branch-shading trees. By far the most important correlate of larval growth was the amount of water consumed with leaf mass (r= 0.94). When variance in water intake was standardized (also largely eliminating parallel variation in proteins), fructose and glucose still had significant positive correlations and proanthocyanidins negative with larval growth on control but not on shade leaves. Concentrations of several phenolic compounds correlated negatively with intake of dry matter and especially water, and different phenolics were important in shaded (gallotannins) and in control (flavonoids) leaves. Our findings strongly suggest that the effects of putatively defensive leaf traits on insect consumption and growth interact with nutritive leaf traits, particularly with water.

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

The quality of available food may affect insect herbivores directly (via growth and survivorship) and/or indirectly (by modifying insect vulnerability to parasitoids and pathogens). We examined the relationship between different phenolic compounds, belonging to various phenolic groups, in Betula pubescens spp. czerepanovii (mountain birch) foliage and the larval performance of the geometrid Epirrita autumnata (autumnal moth). Direct effects on insect performance were described by pupal weight, developmental rate, and survivorship; indirect effects were described by the encapsulation rate of an implant inserted into the insect hemocoel, a commonly used way to describe insect immune defense. We found profound differences in the effects of different phenolic categories: several individual hydrolyzable tannins were associated positively with larval performance but negatively with level of immune defense, whereas flavonoid glycosides were inversely related to larval survival but showed no association with the larvae immune defense.

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

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 Ð from 2 to 57% Ð 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.

Abstract-The surface of birch leaves contains glandular trichomes that secrete exudates containing flavonoid aglycones. We investigated the biological activities of white birch (Betula pubescens) leaf surface exudates against larvae of the autumnal moth, Epirrita autumnata, a common insect pest of birch. We found that tree-specific mortality (up to 100%) of first instar larvae correlated strongly with the tree-specific contents of surface flavonoid aglycones (r s = 0.905) in emerging leaves. We also found that first instars clearly preferred birch buds from which surface exudates had been removed. In addition, the duration of the first instar was shortened by 29%, and the weights and relative growth rates of first instars improved by 8% and 52%, respectively, as a result of removal of the exudates from their leaf diet. The correlation of tree-specific foliar contents of flavonoid aglycones, especially 5-hydroxy-4 ,7-dimethoxyflavanone, with changes in larval performance, suggests that flavonoid aglycones are responsible for the changes observed in first instar larval performance. The results show that chemical characteristics of birch leaves are effective against neonate E. autumnata larvae. However, the removal of leaf surface exudates from fully expanded leaves did not affect the leaf acceptance for the voracious fifth instars. This is probably a result of reduction in contents of flavonoid aglycones compared to those of emerging leaves.