Do Insectivorous Birds use Volatile Organic Compounds from Plants as Olfactory Foraging Cues? Three Experimental Tests

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.     

Do warmer growing seasons ameliorate the recovery of mountain birches after winter moth outbreak?

Subarctic mountain birch (Betula pubescens ssp. czerepanovii) forests in northern Fennoscandia have shown a slight recovery from recent severe defoliation by the winter moth (Operophtera brumata). This development in trees is hypothesized to be a result of ameliorated growing conditions through increased summer temperatures. We examined if accumulated thermal sum affects the ability of mountain birches to tolerate foliage losses. We quantified the number of leaf-bearing short shoots, the emergence of inflorescences and the seasonal height growth of long shoots in both intact and defoliated trees. We also determined the concentrations of carbon and nitrogen in leaves and carbohydrates in roots. Our results show that defoliation constrained the growth of long shoots, as well as the emergence of inflorescences regardless of thermal sum accumulation. However, the number of leaf-bearing short shoots did not differ between intact and defoliated trees. In the both tree groups, the amounts of emerging leaves increased as a response to thermal sum accumulation. Also the leaf carbon concentration increased in defoliated trees at higher thermal sums, whereas it decreased in intact controls. Generally, the mean carbohydrate concentrations were greater in roots of defoliated than intact trees. However, with increased thermal sums, root carbohydrates increased in intact trees but remained the same in defoliated trees. We conclude that thermal sum accumulation does not greatly promote the recovery of mountain birches. Although the damaged trees produced more leaves at warmer growing sites, this did not increase their height growth or carbohydrate gain in roots.     

Delayed induced responses of birch glandular trichomes and leaf surface lipophilic compounds to mechanical defoliation and simulated winter browsing

Changes in morphology and chemistry of leaf surface in response to herbivore damage may increase plant resistance to subsequent herbivore attack; however, there is lack of studies on induced responses of glandular trichomes and their exudates in woody plants and on effects of these changes on herbivores. We studied delayed induced responses in leaf surface traits of five clones of silver birch (Betula pendula Roth) subjected to various types of mechanical defoliation and simulated winter browsing. Glandular trichome density and concentrations of the majority of surface lipophilic compounds increased in trees defoliated during the previous summer. This induced response was systemic, since control branches in branch defoliated trees responded to the treatments similarly to defoliated branches, but differently from control trees. In contrast to defoliation treatments, simulated winter browsing reduced glandular trichome density on the following summer and had fewer effects on individual surface lipophilic compounds. Moreover, constitutive density of glandular trichomes was negatively correlated with induced total amount of lipophilic compounds per trichome, indicating a trade-off between constitutive and induced resistance in silver birch. Induced changes in leaf surface traits had no significant effect on leaf damage by chewers, miners and gall mites, but increased susceptibility of birch trees to aphids. However, leaf damage by chewers, miners and gall mites in defoliated (but not in control) trees was correlated with concentrations of some fatty acids and triterpenoids, although the direction of relationships varied among herbivore species. This indicates that induction of surface lipophilic compounds may influence birch resistance to herbivores. Our study thus demonstrated both specificity of elicitation of induced responses of birch leaf surface traits by different types of damage and specificity of the effects of these responses on different types of herbivores.Electronic Supplementary Material Supplementary material is available in the online version of this article at and is accessible for authorized users.     

Larval parasitism rate increases in herbivore‐damaged trees: a field experiment with cyclic birch feeding moths

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.     

Recovery of a Betula pubescens forest in northern Sweden after severe defoliation by Epirrita autumnata

Abstract. Mountain birch (Betula pubescens ssp. czerepanovii) forest in the Abisko valley of northern Sweden was completely defoliated by Epirrita autumnata caterpillars during an outbreak in 1954–1955. The defoliation resulted in an 80–90% mortality of the leaf‐carrying shoots of birches in 1956 and triggered a rejuvenation of stands. The subsequent regrowth of foliage was studied in two damaged birch stands and in one unattacked stand. The number of leaves approximately doubled in the damaged stands between 1961 and 1987, while the number on the reference plot fluctuated without significant increase. Regrowth started with increased production of long shoots from surviving shoots and basal sprouts. Basal sprouts were a substantial source of new shoots in the recovery of the foliage, especially on the most damaged plot. Trees of seed origin constituted a minor fraction of the regrowth. Initial rapid growth of foliage reduced gradually and the annual leaf production in 1986/1987 was 75% of that of the reference plot. Comparison between the recovery curve and data from the reference plot indicates that the shoot population of the damaged forest will, after more than 30 years, need many more years to reach the assumed size of a mature forest. The degree of rejuvenation varied between stands, with different consequences for future dynamics of E. autumnata populations.     

Delayed induced changes in the biochemical composition of host plant leaves during an insect outbreak

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. Received: 26 January 1998 / Accepted: 10 April 1998     

Movement and disappearance of mountain birch defoliators are influenced by the interactive effects of plant architecture and induced resistance

Abstract. 1. Changes in herbivore movement and feeding behaviour may determine the efficacy of induced plant resistance by affecting the location of damage within the foliage and by modifying the vulnerability of herbivores to predators. 2. Observations of larval feeding sites were used to test whether induced resistance increased the movement of free‐living Epirrita autumnata Borkh. (Lepidoptera, Geometridae) larvae feeding on mountain birch [Betula pubescens ssp. czerepanovii (Orlova) Hämet‐Ahti]. The amount of defoliation at different canopy parts was measured to test the associated changes in the spread of damage within the foliage. 3. The architectural complexity of trees was measured to test its association with the disappearance of larvae from their hosts. The underlying hypothesis was that the architectural traits of the host plant could affect disappearance by influencing the frequency of herbivores encountering predators. 4. Distance between the consecutive feeding positions, the number of leaves damaged, and consumption of long shoot leaves all increased in trees with induced resistance. 5. Disappearance of larvae depended on the architectural complexity of trees. The effect of complexity differed between defoliation treatments, and may depend on the activity and number of predators in relation to the canopy size. 6. Accordingly, this study suggested that the interactive effects of plant architecture, induced resistance, and herbivore behaviour can determine the performance of herbivores on their host plant.     

Low nutritive quality as defence against herbivores: induced responses in birch

Summary The effects of artificial defoliation of birch trees in the previous year on the consumption and utilization of food by a geometrid larva, Epirrita autumnata, were studied in laboratory. The leaves were collected from two sites on a slope of a fell. Defoliation had a significant retarding effect on approximate digestibility, efficiency of conversion of ingested food, relative consumption rate and relative growth rate but not on efficiency of conversion of digested food. The effects were to the same direction with leaves from the two sites, but the response was stronger with leaves from the zone defoliated by Epirrita during mid-sixties. The defensive nature of the response(s) of birch to defoliation is discussed in the light of these results. It is concluded that contrary to the hypothesis of Moran and Hamilton (1980) no increase in the consumption incurred by individual trees could be shown in this system.     

Developmental plasticity in birch leaves: defoliation causes a shift from glandular to nonglandular trichomes

The structures on leaf surfaces, e.g. trichomes, can act as effective antiherbivory mechanisms as chemical repellents. Structural defences usually represent constitutive resistance, but there are also a few cases of inducible morphological defences. We tested whether defoliation may induce changes in trichome production in white birch (Betula pubescens). The studied birches were either 0, 50 or 100% defoliated during the previous or current summer, and we measured the alterations in the production of glandular vs. nonglandular leaf trichomes, developmental instability (fluctuating asymmetry, FA) and leaf and shoot growth. We detected a clear shift from glandular to nonglandular leaf trichomes following previous‐year defoliation but not after current‐year defoliation. Furthermore, the density of nonglandular trichomes around the mid‐vein of leaves increased following previous‐year defoliation but decreased after current‐year defoliation. While leaf and shoot growth showed a distinct decrease in response to defoliation, FA turned out to be less sensitive. Consequently, previous‐year defoliation can induce the production of nonglandular trichomes in birch leaves. Because this response was accompanied by a reduction in glandular trichomes, the present results may suggest a trade‐off between the different trichome types of birch leaves.     

Foliage phenols and nitrogen in relation to growth,insect damage,and ability to recover after defoliation,in the mountain birch Betula pubescens ssp tortuosa

We studied growth of the mountain birch, and the role of foliage phenols, nitrogen, and variance in the timing of bud burst, as potential defensive characters, in Finnish Lapland in 1975–1979. Annual and local variation both in phenol and nitrogen concentration of foliage were significant. Individual trees retained their position in the foliage and nitrogen distribution of the population in successive years, as well as in the order of leaf flush in spring. Growth of twigs, mature leaf size, and ability of trees to recover in the year following artificial defoliation correlated positively with the sum of degree days in the previous growing season. Foliage nitrogen correlated negatively with foliage phenols in within-site comparisons. Twig growth correlated negatively with foliage phenols, particularly in growing seasons following cool summers, but did not correlate with foliage nitrogen. Birches flushing early did not grow more than birches flushing late. Between-site differences in foliage phenol content were mainly determined by abiotic conditions, like temperature and nutrient availability. In a between-site comparison insect chewing marks in leaves correlated positively with foliage phenols as well as with nitrogen; intensity of invertebrate predation presumably explained variable herbivory between the sites. In a within-site comparison trees with the highest foliage phenol content had few herbivores only at the site with the highest average phenol level.     

Consequences of defoliation on phenological interaction between Epirrita autumnata and its host plant, Mountain Birch

1. Defoliation-induced changes in the budbreak phenology of Mountain Birches ( Betula pubescens ssp. tortuosa (Ledeb.) Nyman), and their effects on herbivore performance, were investigated during an outbreak of the spring-feeding geometrid, Epirrita autumnata (Bkh.).
2. Total defoliation (followed by refoliation) by E. autumnata larvae was required to achieve significant delaying of budbreak of Mountain Birch one year after damage.
3. Delayed budbreak has potentially deleterious effects on herbivore performance. However, E. autumnata larvae show phenotypic responses that increase the synchrony between larvae and leaves: high larval density, a prerequisite for severe defoliation, delays egg hatch in the following year.
4. Late hatching larvae performed as well on birches with delayed budbreak due to the previous year's total defoliation as did early hatching larvae on birches with earlier budbreak.     

Effects of warming climate on early-season carbon allocation and height growth of defoliated mountain birches

Tree carbohydrate reserves are usually compromised following insect outbreak, which results in a delay in leaf emergence and a reduction in growth, especially in cold environments. However, in recent times, severe defoliation of subarctic mountain birches (Betula pubescens ssp. czerepanovii) by the winter moth (Operophtera brumata) has not induced such responses. This may be the result of a warming climate stimulating plant primary metabolism. We examined if increasing thermal sum (sum of daily mean temperatures above +5 °C, d.d.) and complete foliage loss affected the concentrations of carbohydrates in sap, juvenile leaves, and fine roots of mountain birches in northern Finland and Norway. The sampling was conducted at the beginning of the growing season, two years after the insect outbreak. We also investigated the morphologic properties of mature leaves and the shoot growth of the trees. Our results showed that the carbohydrate concentrations in leaves and roots (averages 67.8 and 12.5 mg g^?1 DW, respectively) decreased in defoliated trees with increasing thermal sum (>400 d.d.), whereas the response in intact trees was the opposite. The carbohydrates in the sap were unaffected by defoliation or thermal sum accumulation. The leaf area of mature leaves and the height growth of long shoots were greater in trees at warmer sites, irrespective of defoliation. However, defoliation increased the leaf weight per area (SLW: specific leaf weight). We conclude that under warmer growing conditions, low early-season leaf and fine root carbohydrate concentrations of previously defoliated trees cannot be used as indicators of aboveground growth.     

Rapid induced resistance of silver birch affects both innate immunity and performance of gypsy moths: the role of plant chemical defenses

In this study we tested the effects of rapid induced resistance of the silver birch, Betula pendula, on the performance and immune defense of the gypsy moth, Lymantria dispar. We also measured the effects of defoliation on the concentrations of plant secondary metabolites, particularly on phenolics and terpenoids. It was found that severe natural defoliation (by moth larvae) of silver birch led to an increase in lipophilic flavonoids on the leaf surface. The concentration of some simple phenolics and monoterpenes (linalool and geraniol) also increased, while that of several glycosides of quercetin decreased. The female pupal weights and survival rates of moths decreased, and larval development time increased, when the insects fed on defoliated trees. However, the feeding of caterpillars with the leaves of defoliated trees led to an increase in lysozyme-like activity in their hemolymph, with an increase in their ability to encapsulate potential parasites. Our data show that the silver birch deploys a rapid chemical defense against gypsy moth larvae. We suggest that lipophilic flavonoids are important compounds in the direct silver birch defense against L. dispar caterpillars. The increased strength of immune defense of insects exposed to trees that had deployed a rapid induced resistance may be an adaptation of the herbivores to resist the rising density of parasites when host population density is high.     

Effects of defoliation on the saplings of a gap-colonizing neotropical tree

This paper describes a defoliation experiment on saplings of the gap-colonizing pioneer tree Heliocarpus appendiculatus in the tropical rain forest of Los Tuxtlas (Southeast Mexico). Four levels of defoliation (0, 25, 50, and 75% of the leaf area removed) were applied to naturally established plants in a medium-sized forest gap. Records were made of growth (height, diameter, and leaf production) and of whole-plant and leaf survival. Statistical comparisons for all variables showed that plants that had not been defoliated grew better than defoliated ones, but there were no statistically significant differences between the defoliation levels. The risk of death was significantly lower for control plants than for defoliated plants at any level of damage. Leaf survivorship was highly irregular. The most consistent pattern was that the leaves of intact plants always showed higher survivorship, while the most heavily defoliated ones always had the poorest survival. The survivorship pattern of leaves at intermediate defoliation levels was irregular. The results illustrate the lack of a monotonic response to a wide range of defoliation levels, and suggest the potential effect of herbivores as reducers of vegetative growth and survival in pioneer tropical species, and as limiting agents of plant establishment in regenerating forest gaps.     

From plants to birds: higher avian predation rates in trees responding to insect herbivory

BACKGROUND: An understanding of the evolution of potential signals from plants to the predators of their herbivores may provide exciting examples of co-evolution among multiple trophic levels. Understanding the mechanism behind the attraction of predators to plants is crucial to conclusions about co-evolution. For example, insectivorous birds are attracted to herbivore-damaged trees without seeing the herbivores or the defoliated parts, but it is not known whether birds use cues from herbivore-damaged plants with a specific adaptation of plants for this purpose. METHODOLOGY: We examined whether signals from damaged trees attract avian predators in the wild and whether birds could use volatile organic compound (VOC) emissions or net photosynthesis of leaves as cues to detect herbivore-rich trees. We conducted a field experiment with mountain birches (Betula pubescens ssp. czerepanovii), their main herbivore (Epirrita autumnata) and insectivorous birds. Half of the trees had herbivore larvae defoliating trees hidden inside branch bags and half had empty bags as controls. We measured predation rate of birds towards artificial larvae on tree branches, and VOC emissions and net photosynthesis of leaves. PRINCIPAL FINDINGS AND SIGNIFICANCE: The predation rate was higher in the herbivore trees than in the control trees. This confirms that birds use cues from trees to locate insect-rich trees in the wild. The herbivore trees had decreased photosynthesis and elevated emissions of many VOCs, which suggests that birds could use either one, or both, as cues. There was, however, large variation in how the VOC emission correlated with predation rate. Emissions of (E)-DMNT [(E)-4,8-dimethyl-1,3,7-nonatriene], beta-ocimene and linalool were positively correlated with predation rate, while those of highly inducible green leaf volatiles were not. These three VOCs are also involved in the attraction of insect parasitoids and predatory mites to herbivore-damaged plants, which suggests that plants may not have specific adaptations to signal only to birds.     

Comparison of Herbivores and Herbivory in the Canopy and Understory for Two Tropical Tree Species1

The Janzen–Connell model of tropical forest tree diversity predicts that seedlings and young trees growing close to conspecific adults should experience higher levels of damage and mortality from herbivorous insects, with the adult trees acting as either an attractant or source of the herbivores. Previous research in a seasonal forest showed that this pattern of distance‐dependent herbivory occurred in the early wet season during the peak of new leaf production. I hypothesized that distance‐dependent herbivory may occur at this time because the new foliage in the canopy attracts high numbers of herbivores that are limited to feeding on young leaves. As a consequence, seedlings and saplings growing close to these adults are more likely to be discovered and damaged by these herbivores. In the late wet season, when there is little leaf production in the canopy, leaf damage is spread more evenly throughout the forest and distance dependence disappears. I tested three predictions based on this hypothesis: (1) the same species of insect herbivores attack young and adult trees of a given plant species; (2) herbivore densities increase on adult trees during leaf production; and (3) herbivore densities in the understory rise during the course of the wet season. Censuses were conducted on adults and saplings of two tree species, raribea asterolepis and Alseis blackiana. Adults and saplings of both species had largely the same suite of chewing herbivore species. On adults of Q. asterolepis, the density of chewing herbivores increased 6–10 times during leaf production, but there was no increase in herbivore density on adults of A. blackiana. Herbivore densities increased 4.5 times on A. blackiana saplings and 8.9 times on Q. asterolepis saplings during the wet season, but there were no clear trends on the adults of either species. These results suggest that the potential of adult trees as a source of herbivores on saplings depends on the value of new leaves to a tree species' herbivores, which may differ across tree species.     

Influence of previous frost damage on tree growth and insect herbivory of Eucalyptus globulus globulus

The plant stress hypothesis suggests that some herbivores favour stressed plants, whereas the plant vigour hypothesis proposes that other herbivores prefer vigorous plants. The effects of a prior stress, that of frost damage, were examined on the subsequent growth of Eucalyptus globulus globulus and on the response of insect herbivores. Frost damage affected tree growth by reducing new leaf area and increasing specific leaf area (SLA). However, herbivore abundance was not affected by prior frost damage. Two feeding trials using Anoplognathus chloropyrus and Hyalarcta huebneri and a morphometric study of Ctenarytaina eucalypti were conducted to assess the performance of herbivores on trees that had suffered more or less frost damage. Consumption by A. chloropyrus and H. huebneri was unaffected by foliage origin (damaged versus healthy). Hyalarcta huebneri grew faster when fed leaves from previously damaged trees, and C. eucalypti from previously damaged trees were larger than those from healthy trees. Enhanced insect performance on frost damaged plants may have resulted from the high specific leaf area (most likely thinner) leaves. The herbivore abundance data did not support the hypothesis that previously frost damaged plants are preferred by insects. However, increased growth of H. huebneri and larger body size of C. eucalypti on damaged trees indicates that previously stressed trees may produce leaves of higher nutritional value.     

Effects of variable phytochemistry and budbreak phenology on defoliation of aspen during a forest tent caterpillar outbreak

1 The present study assessed the relationship between clonally variable rates of defoliation in trembling aspen (Populus tremuloides Michx.) and two potential resistance traits: defensive chemistry and leaf phenology. 2 In 2001, coincident with a major outbreak of the forest tent caterpillar (Malacosoma disstria Hubner) in the northcentral U.S.A., we monitored defoliation rates, phytochemical composition, and foliar development in 30 clones of trembling aspen. Leaf chemistry was also assessed in re‐flushed leaves and 2 years post‐outbreak. 3 Early in the season, differences in defoliation among clones were substantial but, by mid‐June, all clones were completely defoliated. Leaf nitrogen, condensed tannins, and phenolic glycosides varied among clones but did not relate to defoliation levels. Budbreak phenology differed by 3 weeks among clones and clones that broke bud early or late relative to forest tent caterpillar eclosion experienced reduced rates of defoliation. 4 Defoliation led to increased tannins and slight decreases in phenolic glycoside concentrations in damaged leaf remnants, but to moderately decreased tannins and a six‐fold increase in phenolic glycosides in reflushed leaves. This shift in chemical composition may significantly affect late season herbivores. 5 These results suggest that aspen chemical resistance mechanisms are ineffective during intense episodic eruptions of outbreak folivores such as the forest tent caterpillar. Variable budbreak phenology may lead to differential susceptibility during less intense outbreak years and, at peak forest tent caterpillar population densities, mechanisms affording tolerance are probably more important than chemical defences.     

Effect of defoliation on concentrations of allelochemicals and soluble sugars in leaves of weeping birch (<Emphasis Type="Italic">Betula pendula</Emphasis> roth)

Effect of strong (75%) and complete (100%) artificial defoliation of weeping birch Betula pendula Roth on the dynamics of soluble sugars and phenols—flavonols, catechins, and tannins in leaves of damaged plants was investigated. Within the first 15 days after strong defoliation of birch, no changes were found in leaf contents of flavonol, catechin, and tannin. The concentration of sugars first increased but, on the 10th day after defoliation, it returned to the normal level. One year after strong defoliation, the lead concentrations of catechins and tannins in damaged trees increased, while the concentrations of flavonols and sugars did not differ from that in leaves of control trees. In two years after strong damage, the increased concentration of tannins was retained, while catechins and sugars remained at the control level. One year after complete (100%) artificial defoliation, the leaf concentrations of flavonols and sugars in damaged plants did not differ from that in control plants, while the leaf concentrations of catechins and tannins exceeded those in control plants. Two years after complete damage, the leaves contained an increased amount of tannins, whereas the amounts of catechins, flavonols, and sugars did not differ from the control levels.     

Colonization of a host tree by herbivorous insects under a changing climate

Climate warming has been predicted to increase the abundance of herbivorous insects. Together with concurrent poleward shifts in many insect species this may increase herbivore pressure on plants. However, the manner in which plants at higher latitudes become colonized by herbivorous insects in the future is unknown. We established a translocation experiment using 26 micropropagated silver birch Betula pendula genotypes from six populations originating from 60°N to 67°N, to study the susceptibility of the translocated birches to local herbivores. The birches were planted at three different latitudes in Finland (60°N, 62°N and 67°N). We studied the effect of source population and latitudinal translocation on herbivore density, species richness, and community composition among the genotypes growing in the same environmental conditions in two years; 2011 and 2012. The source population explained the variation in the herbivore density only in 2012, whereas latitudinal translocation did not affect herbivore density. Variation in species richness was not explained by the source population or by the latitudinal translocation. At two of the study sites, the similarity of the herbivore communities among the populations decreased with increasing latitudinal distance of the source populations, possibly because birch populations that grow geographically closer to each other are genetically more similar, and therefore support a more similar composition of the arthropod community. All birch genotypes were colonized by local herbivores, suggesting that as herbivores shift their ranges polewards, they are able to colonize novel host‐plant genotypes. This enables compositional changes in insect communities on their host plants in the future, which in turn, might affect total herbivory and eventually, plant growth.