Delayed inducible resistance against a leaf-chewing insect in four deciduous tree species

Summary Both mechanical damage to mountain birch foliage and rearing of moth larvae on the trees reduced the growth of Epirrita autumnata larvae reared on these trees in the following year. The effects of physical damage and some other cues from insects were additive. On bird cherry the performance of Epirrita larvae was equal on untreated trees and on trees artificially defoliated in the previous year, but larval growth was reduced on previously insect-damaged branches. With mountain ash just physical damage per se reduced the performance of Epirrita larvae. On Salix phylicifolia there were no significant differences in the growth or survival of Epirrita on untreated control bushes and on bushes with partial larval damage during the previous year. Among untreated control trees the growth and survivorship of Epirrita were higher on fast-growing willow and bird cherry than on the slow-growing mountain birch. Mountain birch and mountain ash, the two deciduous tree species adapted to nutrient-poor soils, showed delayed inducible resistance triggered by defoliation (artificial or insect-made). This supports the hypothesis that delayed inducible resistance may be a passive response due to nutrient-stress caused by defoliation. On the other hand, the additional increase in the resistance of mountain birch triggered by specific cues from insects suggests that this response may be an evolved defense against leaf-eating insects.     

The effect of previous defoliation of pole-stage lodgepole pine on plant chemistry,and on the growth and survival of pine beauty moth (Panolis flammea) larvae

Summary A study of the effects of defoliation by insects on the chemistry of lodgepole pine (Pinus contorta), and on the performance of Panolis flammea (Lepidoptera; Noctuidae) larvae, was carried out in a forest in northwest Scotland I year after a severe outbreak of P. flammea had caused extensive defoliation. Larval weight and survival were not significantly different on trees that had experienced different levels of defoliation in 1986. The nitrogen and tannin content of current and previous years' pine needles was not significantly affected by defoliation (although both were slightly greater in the foliage of defoliated trees). Phosphorus content of young pine foliage was lower (but not significantly lower except on one occasion) on heavily defoliated trees. On all sampling occasions, however, the nitrogen: phosphorus ratio was significantly higher on heavily defoliated trees. There were large differences in monoterpene composition of the previous year's shoots associated with defoliation intensity, but these differences had largely disappeared in the new growth. The results are discussed in relation to other studies on the effects of insect damage on plant chemistry and insect performance and in relation to the abundance of P. flammea in Scotland.     

Does tent caterpillar attack reduce the food quality of red alder foliage?

Summary We assayed the quality of red alder trees for western tent caterpillar growth and survival to test the hypothesis that caterpillar feeding stimulates plant defenses in both attacked and adjacent trees. Three years of high tent caterpillar density were necessary before deterioration in foliage quality occurred, and even then only foliage from trees which were almost completely defoliated in the current year reduced the growth of caterpillars. Both tent size and mean egg mass size increased after the second year of high density which indicates that good conditions still existed for tent caterpillars after 2 to 3 years of heavy feeding.Egg masses which were moved to areas where trees had not recently supported a high caterpillar population produced significantly smaller tents than endemic controls in 1982. Therefore the small tent and egg mass size of the high density population in 1982 was inherent to the insects rather than modified by food source. In 1983 the tents from introduced egg masses were as large as naturally occurring tents.If lightly attacked trees within areas of high caterpillar density are better defended against insect attack, this does not show up in their ability to support caterpillar growth and survival. We found no evidence to support the hypothesis that trees communicate insect attack and stimulate chemical defenses in adjacent trees. Reduced foliage quality seems to be a result of extensive insect damage rather than a defense against insect damage.     

The inhibition of phenolic biosynthesis in damaged and undamaged birch foliage and its effect on insect herbivores

Summary 1. The leaves of Betula pendula Roth trees were damaged artificially, or by insect-grazing. Both induced an increase in phenolic levels in damaged leaves, larger in the case of insect attack.-2. Some of the damaged trees were sprayed with an inhibitor of phenolic biosynthesis, (aminoxy) acetic acid, which led to a reduction in phenolic levels in both undamaged and damaged leaves. Hence both the effects of damage per se and damage-induced changes in foliage phenolic levels on insect feeding preference could be examined using this technique.-3. Herbivore feeding preferences were assessed in the laboratory by comparing damaged and undamaged leaves, with or without phenolic inhibition, using caterpillars of a natural birch feeder, Apocheima pilosaria D. & S. (Lepidoptera: Geometridae) and a non-birch feeder, Spodoptera littoralis Boisduval (Lepidoptera: Noctuidae). Neither species showed any significant preferences and appeared indifferent to damage, irrespective of whether the trees had their damage-induced phenolic synthesis blocked.-4. The implications of these results for induced defense theory are discussed.     

Recycling of nitrogen in herbivore feces: plant recovery,herbivore assimilation,soil retention,and leaching losses

Herbivores directly and indirectly affect ecosystem functioning in forests. Feces deposition is a direct effect that supplies ephemeral N pulses to soils. Herbivore-mediated changes in plant N allocation and uptake are indirect effects that can also influence soil N availability. These effects may interact if defoliation influences the ability of plants to recover fecal N, and this may affect subsequent generations of herbivores. We added ¹⁵N-enriched insect feces (frass) to a series of replicated red oak, Quercus rubra, mesocosms that had been damaged experimentally and then followed the frass N over the course of 2 years. In the first season, some frass N was mineralized in the soil and leached in organic form from the mesocosms within 1 week of deposition. Within 1 month, frass N had been acquired by the oaks and enriched the foliage; late-season herbivores assimilated the frass N within the same growing season. In the second season, herbivore damage from the previous year lowered total leaf N contents and ¹⁵N recovered in the foliage. A subsequent cohort of early-season herbivores fed on this foliage consequently derived less of their N from the previous year’s frass, and feral leaf rollers colonized fewer of these saplings. The 0- to 5-cm soil fraction was the largest N sink measured, and 42% of the frass N was recovered in the soil. The results demonstrate that: (1) some frass N can be recycled rapidly into foliage and assimilated by successive cohorts of herbivore within the same season; (2) damage can affect N allocation in the following year’s foliage, influencing N availability to and host selection by herbivores; and (3) leaching losses occur soon after deposition but are buffered by soil pools, which are the largest sinks for frass N.     

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     

The influence of ants on the survival of mountain birches during an Oporinia autumnata (Lep., Geometridae) outbreak

Summary The present study reports the influence of ants on the survival of mountain birch (Betula pubescens ssp. tortuosa) during an Oporinia autumnata (Lep., Geometridae) outbreak. Undamaged green islands with a radius of 15–20 m were observed around Formica aquilonia mounds in a defoliated area in Finnish Lapland. The herbivore densities and grazing pressure were shown to be low within this radius. The foraging strategy of ants and their influence on birch growth are discussed.     

Defoliation increases carbon limitation in ectomycorrhizal symbiosis of <Emphasis Type="Italic">Betula pubescens</Emphasis>

Boreal forest trees are highly dependent on root-colonizing mycorrhizal fungi. Since the maintenance of mycorrhizal symbiosis implies a significant carbon cost for the host plant, the loss of photosynthetic leaf area due to herbivory is expected to reduce the host investment in mycorrhizae. We tested this hypothesis in a common garden experiment by exposing ectomycorrhizal white birch (Betula pubescens Ehrh.) seedlings to simulated insect defoliation of 50 or 100% intensity during either the previous or the current summer or repeatedly during both seasons before harvest. The shoot and root growth of the seedlings were distinctly reduced by both 100% defoliation and repeated 50% defoliation, and they were more strongly affected by previous-year than current-year defoliation. The root to shoot ratio significantly decreased after 100% defoliation, indicating reduced proportional allocation to the roots. Ergosterol concentration (i.e. fungal biomass) in the fine roots decreased by 100% defoliation conducted either in the year of harvest or in both years. No such decrease occurred following the 100% defoliation conducted in the previous year, indicating the importance of current photosynthates for fungal symbionts. The trend was similar in the colonization percentage of thick-mantled mycorrhizae in the roots, the most marked decline occurring in the repeatedly defoliated seedlings. The present results thus support the prediction that the plant investment in ectomycorrhizae may decline as a response to foliage loss. Moreover, the colonization percentage of thick-mantled mycorrhizae correlated positively with the ratio of leaf to heterotrophic plant biomass in the defoliated birch seedlings, but not in the control ones. This tends to indicate a stronger carbon limitation of ectomycorrhizal colonization in defoliated seedlings.     

Intraspecific variation in food plant favourability to phytophagous insects: psyllids on Eucalyptus blakelyi M.

Abstract.

• 1 A combination of variables reflecting leaf quality provided the best predictive model explaining differences in population trend of Glycaspis spp. among individuals of the food plant Eucalyptus blakelyi, measured under field conditions on an egg to egg basis. The equation incorporated the product of: (1) proportion of foliage in each age class; (2) probability of foliage escape from herbivory; (3) preference for the foliage class measured as the proportion of eggs oviposited on it by the female; these variables being summed over the young and mature age classes.
• 2 In addition, significant and consistent differences in insect survivorship among trees were demonstrated by transfer experiments. These experiments established that young foliage was preferable to mature foliage for first instar survival. Differences in insect survivorship also showed a consistent pattern among the study trees. The most probable explanation for this consistent pattern was the influence of the trees on the insects, particularly physiological or biochemical differences among trees. It was not, however, possible to identify the cause of the differences.

     

Insect outbreaks alter nutrient dynamics in a southern African savanna: patchy defoliation of Colophospermum mopane savanna by Imbrasia belina larvae

Invertebrate herbivore outbreaks have important impacts on system biogeochemical cycling, but these effects have been poorly documented in African savanna ecosystems. In semi‐arid African savannas, outbreaks of the lepidopteran Imbrasia belina (mopane worm) affect discrete patches of the dominant Colophospermum mopane trees; larvae may completely defoliate trees for up to six weeks during each of the early and late growing seasons. We studied the impact of mopane worm outbreaks on the availability of nitrogen (N), phosphorus (P), and potassium (K) within mopane savanna by quantifying major nutrient pools in defoliated and non‐defoliated savanna patches, including leaves, leaf litter, worm frass, and the soil beneath trees. Within an outbreak area, approximately 44 percent of trees were infested, supporting ~29,000 worms/ha, leading to ~640 kg/ha dry weight frass deposition at 1.4 g of frass/day‐individual (fourth or fifth instar), compared with an average 1645 kg/ha dry weight of leaf on trees most of which should be deposited by litterfall at the end of the growing season. Frass had twofold higher P, 10 percent higher K, but equivalent N content than litter. Taking frass and litter deposition together, the N, P, and K contents added due to the outbreak event at our study site were 0.88, 5.8, and 2.8 times those measured in non‐outbreak patches, a pattern which was reflected in the nutrient contents of soil surfaces beneath defoliated trees. Invertebrate herbivory appears to be an important driver for mopane savanna but has been largely neglected.     

Dieback of rural eucalypts: Response of foliar dietary quality and herbivory to defoliation

Rural dieback of Eucalyptus blakelyi trees growing on pastoral properties near Canberra is associated with chronic defoliation by insects. In order to test the effect of defoliation on subsequent herbivory, I artificially defoliated three healthy trees by clipping their terminal branchlets. The foliage that regrew on the clipped trees was nutritionally superior to the foliage it replaced, and was much more heavily damaged by grazing insects. There was a transient increase in the tannin content of the regrowth foliage, but this was apparently ineffective in defending it from subsequent herbivory. Compared with the foliage on nearby E. blakelyi trees that also produced major flushes of leaf growth during the same period, the regrowth on the clipped trees had enhanced dietary qualities and suffered more insect damage. Leaf age contributed to many of the differences in dietary quality, but when adjustments were made for the effects of leaf age the same trends remained. Five of the nearby trees were suffering from the chronic insect grazing associated with rural dieback, and the other five appeared healthy. The dietary quality of regrowth foliage on the clipped trees was qualitatively more similar to that of foliage on the dieback trees. Thus the chronic herbivory associated with rural dieback may be partly self-perpetuating, given this positive feedback between defoliation and dietary quality, and an apparent absence of other effective controls on insect populations.     

Green Islands — nutrition not predation —an alternative hypothesis

Summary Mountain birch trees are said to survive as green islands around nests of red ants in Finnish Lapland because the ants kill larvae which would defoliate trees during outbreaks of the moth O. autumnata. An alternative hypothesis says that because the ants will concentrate soil nutrients (and possibly ameliorate soil moisture and temperature) in and around their nests, they provide a more favourable site for trees growing nearby. These trees are therefore less stressed and a poorer source of food for defoliators at times of outbreaks. Few if any young O. autumnata larvae survive on the trees which then survive in green islands around ant nests.     

The feeding behaviour of a sit-and wait-predator,Ranatra dispar (Heteroptera: Nepidae): optimal foraging and feeding dynamics

Summary Nitrogen and phosphorus flow in litterfall and throughfall were studied in two California Quercus species (the evergreen Q. agrifolia and deciduous Q. lobata) before, during, and after an outbreak of the California oak moth, Phryganidia californica. All of the foliage of both oak species was removed by the herbivore during the course of this outbreak. During the outbreak, total N and P flow to the ground more than doubled from Q. agrifolia and increased to a lesser extent from Q. lobata over the previous year. The composition of the litter during the outbreak year shifted so that in Q. agrifolia, almost 70% of the total N and P flow to the ground moved through frass and insect remains, while in Q. lobata, approximately 60% of the N and 40% of the P moved through frass and insect remains. Short-term leaching experiments showed that nitrogen was far more rapidly lost from Phryganidia frass than from leaf litter of either species. These results and the relative frequency of Phryganidia outbreaks suggest that this herbivore has significant effects on the nutrient cycling beneath these trees.     

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.     

The evolution of offspring size and number: a test of the Smith-Fretwell model in three species of crickets

Most models of parental investment in offspring assume a trade-off between propagule size and number, and an increasing concave down function relating offspring fitness to propagule size. In this study, we test these two fundamental assumptions, using three closely related species of crickets, Gryllus firmus, G. veletis, and G. pennsylvanicus. Egg weight, 35-day fecundity and 35-day egg biomass were estimated in a population of each species, and the relationships between these reproductive traits and date of egg laying and body size were estimated. The relationships between egg weight and offspring survival were also sought for eggs buried at different depths, soil moistures, and soil types (G. firmus and G. veletis), as well as in the field (G. pennsylvanicus). A trade-off between egg weight and 35-day fecundity was revealed in a multivariate analysis taking into account among-species variation in egg weight and body size. Independent of the environmental conditions affecting the eggs, a positive correlation existed between the number of larvae that emerged from the soil and propagule weight in each species. Therefore, these findings provide partial support for the assumptions considered in the models mentioned above. A single optimal egg size was favored in two out of the three sets of conditions in which the functions relating egg weight to larval survival could be derived. The conditions encountered by the eggs, however, influenced the average survival of the larvae, as well as the shape of the relationship between egg weight and offspring survival. This suggests that cricket eggs frequently face heterogeneous environments with respect to egg and hatchling survival; the implication of habitat heterogeneity on the evolution of an optimal egg size is considered. The relationships between the reproductive components and female age and size, as well as between egg size and variation in cricket life-history, are discussed in an ecological and evolutionary context.     

Numerical and functional responses of breeding passerine species to mass occurrence of geometrid caterpillars in a subalpine birch forest: a 30-year study

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

Leaf production of an overstorey rainforest tree and its effects on the temporal distribution of associated insect herbivores

Summary The host phenology and temporal distribution of insect herbivores associated with a rainforest canopy tree, Argyrodendron actinophyllum (Sterculiaceae), were monitored for 3 and 2 years respectively in an Australian subtropical rain forest near Brisbane. Leaf production of this evergreen species was synchronous among individuals, occurring from October to February, but differed markedly between years for some fast-growing trees. Most herbivore species fed selectively upon young leaves and, consequently, occurred only during windows of young foliage availability, with the exception of some mesophyll-feeders that preferred mature foliage. Fast-growing trees sustained high herbivore activity during periods of intense leaf production but some slow-growing trees, whose availability of young foliage appeared to be more constant and predictable, also supported high herbivore activity in the long term. Host suitability of particular trees during study years was dependent upon the intensity of leaf production and other unidentified factors, and was different for generalist and specialist herbiovores. Certain specialist species, such as psyllids, were able to overcome the changing availability of young foliage between years on vigorous A. actinophyllum trees by feeding upon both vegetative and reproductive meristems. Most of the data suggest that availability of young foliage is an important factor in this insect-plant system.     

Refoliation of deciduous canopy trees following severe insect defoliation: comparison of <Emphasis Type="Italic">Fagus crenata</Emphasis> and <Emphasis Type="Italic">Quercus crispula</Emphasis>

Deciduous trees can survive severe defoliation by herbivores and often refoliate in the same season. Refoliation following severe defoliation represents compensatory regrowth to recover foliage biomass. Although the relationship between defoliation intensity and degree of refoliation at the individual level has been quantified following artificial defoliation for saplings and small trees, no study has examined the relationship for canopy trees and interspecific differences in this relationship. In this study, defoliation by gypsy moths in an outbreak year and subsequent refoliation were visually surveyed for canopy trees of Fagus crenata (n?=?80) and Quercus crispula (n?=?113) in central Japan. Defoliation and refoliation estimates were scored in 10% classes as the ratio to foliage present before defoliation. The degree of refoliation and the proportion of refoliated trees were high in severely defoliated trees. For 60 and 100% defoliated trees, respective refoliations were 2 and 66% for F. crenata, and 37 and 88% for Q. crispula. All of the 90 and 100% defoliated trees refoliated. These results indicate that severely defoliated trees show an increased need for refoliation to maintain metabolism. Beta regression analysis showed that Q. crispula possessed higher refoliation capability than F. crenata. This is likely associated with the relatively large storage reserves and recurrent growth flush pattern of oak species, which are strong characteristics of oaks and adaptive for response to herbivory and catastrophic disturbances. Interspecific differences in refoliation capability may exert differential effects on forest ecosystem processes, such as influencing the growth of understory species.     

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.     

Biology of the Eucalyptus leaf beetle Paropsisterna selmani (Chrysomelidae: Paropsini): a new pest of Eucalyptus species (Myrtaceae) in Ireland

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