Performance of the green spruce aphid, Elatobium abietinum (Walker) on previously defoliated Sitka spruce

Abstract 1 The green spruce aphid, Elatobium abietinum, is an important defoliator of Sitka spruce in the U.K. However, it is usual for years in which high E. abietinum populations occur to be followed by a year with low aphid densities. The possibility that the performance of E. abietinum is reduced on previously infested Sitka spruce, and that this is the cause of year‐to‐year fluctuations in population density, was investigated by comparing population development and the growth rate of individual aphids on experimentally defoliated trees. 2 Separate experiments were performed to determine whether aphid performance was reduced either in the autumn immediately after defoliation in the spring, or was reduced in the spring of the next year. Different rates of initial defoliation on trees used to test aphid performance were created by artificially infesting the trees with aphids in the spring before the experiments, and varying the time of infestation. 3 Population development and the mean relative growth rate (MRGR) of individual aphids on previously defoliated and undefoliated Sitka spruce did not differ significantly in the spring of the next year. No differences were observed in the nutrient content of the 1‐year‐old needles of previously defoliated or undefoliated trees at this time. 4 In the autumn and winter immediately after spring defoliation, aphid MRGR was significantly higher on trees that had been heavily defoliated earlier in the season compared with trees that had been lightly defoliated. However, the difference in MRGR decreased over the winter period. Nitrogen, phosphorous and potassium concentrations were 9.4–12.2% higher, at the beginning of the autumn, in the current year needles of heavily defoliated trees than in the current year needles of lightly defoliated trees. 5 The experiments indicate that high populations of E. abietinum in the spring do not induce any defensive mechanisms in Sitka spruce that adversely affect subsequent generations of the aphid. By contrast, the results suggest that high spring densities of the aphid improve the nutritional quality of the current year's foliage for autumn generations.     

Growth and mortality of trembling aspen (Populus tremuloides) in response to artificial defoliation

To simulate the effects of forest tent caterpillar (FTC) defoliation on trembling aspen growth and mortality, an artificial defoliation experiment was performed over three years in young aspen stands of northwestern Quebec. Defoliation plots of 15 × 15 m were established on three sites, together with associated control stands of pure trembling aspen. In 2007, root collar diameters were measured and positions of all trees were mapped prior defoliation. Severe FTC defoliation was simulated for three successive years (2007–2009) by manually removing all leaves from all but 7–10% of the trees present in the defoliation plots. Yearly surveys of growth and mortality were conducted until 2010 to evaluate defoliation effects on defoliated as well as surrounding undefoliated trees. In absence of other factors, growth and mortality of trembling aspen decreased and increased, respectively, after defoliation. Our study further revealed that small diameter trees died after one year of artificial defoliation, while larger-diameter trees died after repeated defoliations. Distributions of tree mortality tended to be aggregated at small scales (<5 m), corroborating gap patterns observed in mature stands following FTC outbreaks. This experiment revealed that trembling aspen mortality can be directly attributed solely to defoliation. Repeated defoliations during FTC outbreaks have the potential to profoundly modify stand productivity and structure by reducing tree growth and increasing tree mortality in the absence of predisposing factors.     

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.     

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     

Larch- and pine-feeding host races of the larch bud moth (Zeiraphera diniana) have cyclic and synchronous population fluctuations

Population cycles of many forest-defoliating insects often show synchronous fluctuations at both intra-specific (spatial synchrony) and inter-specific levels. However, population dynamics of different host-associated biotypes of the same species, such as those of the larch budmoth (LBM), Zeiraphera diniana (Lepidoptera: Tortricidae), have never been compared. This species causes extensive defoliation of larch forests every 8 to 9 years in the Alps, but it consists of two genetically-differentiated host races, the first one developing on European larch, Larix decidua , and the other one developing on Swiss stone pine, Pinus cembra . The dynamics of Zeiraphera populations have been extensively studied on larch, whereas little is known about larval density and possible population fluctuations on sympatric pines. A larval census on Swiss stone pine was conducted in the Swiss Alps intermittently between 1958 to 2004 and in the French Alps from 1992 to 2004. Population density of Zeiraphera on pine varied up to 5000-fold and showed periodic oscillations, with five peaks in Switzerland and one in France. Because the feeding activity of the pine race is restricted to the elongating shoot of the current year, no conspicuous defoliation of pine trees was noted during years of high larval densities. Zeiraphera populations on pine oscillated in significant synchrony with larch-associated populations, and peak densities were observed either the same year or shifted by±one year. Our results did not allow any explanation for cyclic fluctuations of LBM on pine, but the synchrony with the larch race's cycle suggests that studies on genetics as well as on parasitism should be intensified.     

Possum (

To assess the effect of possum browse on plant growth, an index of the amount of foliage on about 50 trees of Fuchsia excorticata and the number of trees that died or were completely defoliated was measured at five sites in South Westland over 5 years. This index was compared to possum density indices taken at each site each year. At one site, possums were reduced from a high density about 6 months before the final measurement. The degree of defoliation of fuchsia was significantly related to the density of possums at each site. Possums defoliate and kill fuchsia within two years once the catch-rate index of their density exceeds about 25%. However, this impact can be halted and quickly reversed by reducing possum density.     

A lagged, density-dependent relationship between jack pine budworm Choristoneura pinus pinus and its host tree Pinus banksiana

Abstract. 1. Survival of newly emerged jack pine budworm Choristoneura pinus pinus is related to the density of available pollen cones (microsporangiate strobili) produced by its host tree, jack pine Pinus banksiana. 2. A 7‐year time series of observations from a plot network in Ontario, Canada, compared the propensity of jack pine to produce pollen cones, τ, on trees that were either defoliated or undisturbed by the jack pine budworm. 3. Non‐defoliated jack pine trees have a high propensity to produce pollen cones. More than one‐third of these trees produced pollen cones in every year of the series. Propensity varied significantly among plots and trees. Temporal patterns in propensity were also highly variable but within a plot propensity was often autocorrelated in time. 4. Defoliation by the jack pine budworm was associated with forest plots composed of the oldest and the largest trees and with the fewest trees per hectare. Within a plot, outbreaks lasted 3 or 4 years although individual trees were only defoliated in 1 or 2 years. 5. The propensity to produce pollen cones in jack pine was reduced in the years after defoliation. The most pronounced reductions in propensity occurred where defoliation was most severe. 6. The reduction in propensity to produce pollen cones resulting from previous defoliation, coupled with the dependence of jack pine budworm survival on the availability of pollen cones, induces a lagged, negative feedback between the density of the consumer and that of its resource. 7. The lagged, density‐dependent relationship between jack pine budworm and its jack pine host contributes to oscillatory dynamics of the jack pine budworm. Comparison of the outbreak behaviour of jack pine budworm with that of the closely related eastern spruce budworm C. fumiferana suggests that differences in the strength of the host‐plant interaction may account for differences in the relative frequency of outbreaks in the respective systems.     

Defoliation of Sitka spruce by the European spruce sawfly, Gilpinia hercyniae (Hartig): a retrospective analysis using the needle trace method

Abstract 1 During 1968–74, an outbreak of the European spruce sawfly Gilpinia hercyniae, defoliated many stands of Sitka spruce in commercial forests in mid‐Wales. The needle trace method was used to determine retrospectively the temporal pattern and intensity of defoliation in a stand of Sitka spruce in Hafren Forest that had been damaged severely at the time of the sawfly outbreak. 2 An initial calibration experiment, designed to test the reliability of the needle trace method when applied to Sitka spruce, indicated that artificially induced defoliation of up to 75% was detected by the technique with an accuracy of ±7%. Higher rates of defoliation were underestimated by up to 17%. 3 For the main sample of trees, the needle trace method demonstrated that retention of needle sets (average needle retention) was reduced by up to 33–38% over a 10‐year period coincident with the G. hercyniae outbreak and the years immediately afterward when the canopy was recovering. 4 Analysis of needle loss within separate needle cohorts and age‐classes revealed that 1‐year‐old needles were the most severely defoliated. The density of 1‐year‐old needles was reduced by 51–78% in 1970–73. 5 Defoliation at the time of the G. hercyniae outbreak was associated with reductions in annual height, radial, and volume increments of 24–49%, 30–59% and 32–56%, respectively. Radial and volume increments suffered their greatest reductions 1 year later than height increments. 6 The study demonstrates that the needle trace method can be applied successfully to Sitka spruce to quantify defoliation caused by an insect outbreak occurring many years previously, and that the technique can provide data on needle loss that is valuable for interpreting reductions in tree growth.     

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.     

The effect of defoliation intensity and history on photosynthesis, growth and carbon reserves of two conifers with contrasting leaf lifespans and growth habits

The effects of partial defoliation on photosynthesis, whole-seedling carbon allocation, partitioning and growth were studied for two species with contrasting foliar traits. Field-grown seedlings of deciduous Japanese larch ( Larix leptolepis ) and evergreen red pine ( Pinus resinosa ) were defoliated by hand in early summer for 2 consecutive years. In the first year (1990), seedlings were defoliated by removing the distal 0, 25, 50 or 75% of each needle. In the second year (1991), seedlings were defoliated either 0 or 50%, regardless of previous defoliation treatments. Defoliation had little effect on photosynthesis and starch concentration in whole seedlings of either species in the first year. In the second year, photosynthesis increased in both species in response to the 1991 defoliation treatment, and in red pine also increased in response to the 1990 defoliation treatment. Further, in 1991 both larch and pine had decreased whole-seedling total non-structural carbohydrate concentrations in all seedlings that were defoliated at least once over the 2-yr period. This decrease was noted mostly in the starch component of the non-structural carbohydrates, and was similar in both species. In 1991, biomass was similarly decreased in both species in response to 1991 defoliation. Both species showed overcompensation in total and component biomass in seedlings defoliated by 25% in 1990. Overall, the results do not support the widely held belief that evergreen trees are substantially more affected than deciduous trees by defoliation.     

Artificial Defoliation Induces Trichome Production in the Tropical Shrub Cnidoscolus aconitifolius (Euphorbiaceae)1

Induction of plant defenses and their spatial variability are key subjects in the field of ecology and evolution of defensive traits in plants. Nevertheless, induction has been more commonly studied under controlled environments, ignoring other factors that might influence this process in natural settings. The main goal of this study was to determine if artificial defoliation induces trichome production in three natural populations of the tropical shrub Cnidoscolus aconitifolius. First, we performed trichome counts for each population before imposing artificial defoliation to assess differences in trichome loads between populations. Trichome densities (trichomes/cm²) were quantified for leaf blades, petioles, and flower stalks. To determine if defoliation induced trichome production, three defoliation treatments (0% leaves defoliated or controls, 50% of total leaves defoliated, and 100% defoliation) were applied once at the beginning of the reproductive season. Trichome counts were performed on each structure every ～20 d during a 3‐mo period after the application of treatments. Trichome counts showed significant differences in trichome densities between populations for all three structures. In turn, artificial defoliation increased trichome density. Significant differences among treatments were found for trichome densities on leaf blades and petioles. In both these structures, the 100 percent defoliation treatment differed significantly from control plants, presenting higher trichome densities. In addition, the treatment × population interaction was not significant for leaf blades and petioles, indicating that induction is a generalized response in this species, at least at the study sites. These results indicate that trichomes in C. aconitifolius are inducible due to defoliation.     

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.     

Enriched atmospheric CO2 and defoliation: effects on tree chemistry and insect performance

We examined the effects of CO₂ and defoliation on tree chemistry and performance of the forest tent caterpillar, Malacosoma disstria. Quaking aspen (Populus tremuloides) and sugar maple (Acer saccharum) trees were grown in open-top chambers under ambient or elevated concentrations of CO₂. During the second year of growth, half of the trees were exposed to free-feeding forest tent caterpillars, while the remaining trees served as nondefoliated controls. Foliage was collected weekly for phytochemical analysis. Insect performance was evaluated on foliage from each of the treatments. At the sampling date coincident with insect bioassays, levels of foliar nitrogen and starch were lower and higher, respectively, in high CO₂ foliage, and this trend persisted throughout the study. CO₂-mediated increases in secondary compounds were observed for condensed tannins in aspen and gallotannins in maple. Defoliation reduced levels of water and nitrogen in aspen but had no effect on primary metabolites in maple. Similarly, defoliation induced accumulations of secondary compounds in aspen but not in maple. Larvae fed foliage from the enriched CO₂ or defoliated treatments exhibited reduced growth and food processing efficiencies, relative to larvae on ambient CO₂ or nondefoliated diets, but the patterns were host species-specific. Overall, CO₂ and defoliation appeared to exert independent effects on foliar chemistry and forest tent caterpillar performance.     

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.     

Extreme defoliation reduces tree growth but not C and N storage in a winter-deciduous species

Background and Aims There is a growing concern about how forests will respond to increased herbivory associated with climate change. Carbon (C) and nitrogen (N) limitation are hypothesized to cause decreasing growth after defoliation, and eventually mortality. This study examines the effects of a natural and massive defoliation by an insect on mature trees’ C and N storage, which have rarely been studied together, particularly in winter-deciduous species.Methods Survival, growth rate, carbon [C, as non-structural carbohydrate (NSC) concentration] and nitrogen (N) storage, defences (tannins and total polyphenols), and re-foliation traits were examined in naturally defoliated and non-defoliated adult trees of the winter-deciduous temperate species Nothofagus pumilio 1 and 2 years after a massive and complete defoliation caused by the caterpillar of Ormiscodes amphimone (Saturniidae) during summer 2009 in Patagonia.Key Results Defoliated trees did not die but grew significantly less than non-defoliated trees for at least 2 years after defoliation. One year after defoliation, defoliated trees had similar NSC and N concentrations in woody tissues, higher polyphenol concentrations and lower re-foliation than non-defoliated trees. In the second year, however, NSC concentrations in branches were significantly higher in defoliated trees while differences in polyphenols and re-foliation disappeared and decreased, respectively.Conclusions The significant reduction in growth following defoliation was not caused by insufficient C or N availability, as frequently assumed; instead, it was probably due to growth limitations due to factors other than C or N, or to preventative C allocation to storage. This study shows an integrative approach to evaluating plant growth limitations in response to disturbance, by examining major resources other than C (e.g. N), and other C sinks besides storage and growth (e.g. defences and re-foliation).     

Resistance in oak (Quercus spp.) to defoliation by Tortrix viridana L. in Roudsea Wood National Nature Reserve

Resistance in oak ( Quercus spp.) to defoliation by caterpillars of Tortrix viridana L. was studied in an oak wood and a mixed wood in Roudsea Wood National Nature Reserve. When defoliation is severe in the oak wood a few trees there and most of the oaks in the mixed wood retain relatively undamaged canopies. Their resistance was found to be phenological. The resistant trees in the oak wood flush either later or earlier than those that become defoliated; those in the mixed wood flush later. On late-flushing trees the buds open too late for newly hatched larvae to penetrate them, the resultant mortality reducing the larval population below the level at which damage is severe. The early-flushing trees carry as many larvae per leaf cluster as non-resistant trees but do not show severe defoliation because they produce more foliage and 'grow away' from the attack.     

Attack dynamics of the pine shoot beetle, Tomicus piniperda (Col.; Scolytinae) in Scots pine stands defoliated by Bupalus piniaria (Lep.; Geometridae)

Abstract 1 After a 1‐year, extensive pine looper (Bupalus piniaria) outbreak, plots were laid out to study tree susceptibility to attack, and performance of Tomicus piniperda in pine trees suffering from varying levels of defoliation. 2 Tomicus piniperda was the dominating stem‐attacking species among the primary stem colonizers, and 82% of all trees that died had been colonized by T. piniperda. 3 Beetle attacks primarily struck severely defoliated trees, i.e. trees that suffered from 90% to 100% defoliation. 4 Beetle attacks peaked in the second year after cessation of the outbreak, and suppressed trees were both more frequently attacked and more susceptible to beetle attack than intermediate and dominant trees. 5 Trees surviving beetle attacks carried more foliage than trees that did not survive the attacks. 6 A single year of severe defoliation is enough to render pine trees susceptible to secondary pests, such as T. piniperda.     

Crown defoliation decreases reproduction and wood growth in a marginal European beech population

Background and AimsAbiotic and biotic stresses related to climate change have been associated with increased crown defoliation, decreased growth and a higher risk of mortality in many forest tree species, but the impact of stresses on tree reproduction and forest regeneration remains understudied. At the dry, warm margin of species distributions, flowering, pollination and seed maturation are expected to be affected by drought, late frost and other stresses, eventually resulting in reproduction failure. Moreover, inter-individual variation in reproductive performance versus other performance traits (growth, survival) could have important consequences for population dynamics. This study investigated the relationships among individual crown defoliation, growth and reproduction in a drought-prone population of European beech, Fagus sylvatica.MethodsWe used a spatially explicit mating model and marker-based parentage analyses to estimate effective female and male fecundities of 432 reproductive trees, which were also monitored for basal area increment and crown defoliation over 9 years.Key ResultsFemale and male fecundities varied markedly between individuals, more than did growth. Both female fecundity and growth decreased with increasing crown defoliation and competition, and increased with size. Moreover, the negative effect of defoliation on female fecundity was size-dependent, with a slower decline in female fecundity with increasing defoliation for the large individuals. Finally, a trade-off between growth and female fecundity was observed in response to defoliation: some large trees maintained significant female fecundity at the expense of reduced growth in response to defoliation, while some other defoliated trees maintained high growth at the expense of reduced female fecundity.ConclusionsOur results suggest that, while decreasing their growth, some large defoliated trees still contribute to reproduction through seed production and pollination. This non-coordinated decline of growth and fecundity at individual level in response to stress may compromise the evolution of stress-resistance traits at population level, and increase forest tree vulnerability.     

Anti-herbivore strategies in mountain birch at the tree-line

Birch forest on the slope of the Jesnalvaara fell forms a gradient ranging from 7–8 m high trees (at 75 m a.s.l.) to birch shrubs less than 1 m high at the top of the fell (330 m a.s.l.). Upper parts of the forest (above 220 m) were defoliated by Oporinia autumnata (Lep., Geometridae) about ten years ago and, with the exception of the top, only a fraction of birches have recovered. Climatic conditions explain the limitation of the damaged area – Oporinia eggs were killed during cold winters. Predators may also limit the damage. Defensive mechanisms in the birch leaves – especially a low nitrogen content and high concentrations of phenols inhibiting trypsin – can slow down an increase in herbivore density by reducing their reproductive capacity. Birch leaves from the lower limit of the damaged area are least suitable for the growth of herbivores. They have the highest concentration of phenols, too. Birches at the foot of Jesnalvaara are able to increase leaf phenols after mechanical damage of nearby leaves. This is enough to retard the growth of several herbivore species. Ability to recover after defoliation is an important part of the anti-herbivore strategy of the birch. This ability is higher in shrub birches than in tall trees and better after a warm than a cold summer. Birch leaves in the year following defoliation are less usable for herbivores and cause increased mortality and lowered reproduction. These properties are the weakest in birches at the foot of Jesnalvaara where the probability of being defoliated is also lowest.     

Effects of previous defoliation on pine looper larval performance

1 During outbreaks of the pine looper, Bupalus piniarius, its host, Pinus sylvestris, is severely defoliated. The larvae of this geometrid normally feed on mature needles. However, because trees are totally defoliated during outbreaks, the next generation is forced to feed on current-year needles. 2 Bupalus piniarius larvae feeding on previously defoliated trees may show lower performance either because of induced resistance or because larvae have to feed on needles not normally fed upon (current instead of mature). 3 These hypotheses were tested in an experiment where larvae were reared on (i) shoots naturally defoliated the previous year, and thus, bearing only current-year needles, (ii) non-defoliated shoots where larvae had access only to current-year needles, and (iii) control shoots with access to both current and mature needles. 4 There was no support for the induction hypothesis. Survival was lower on naturally defoliated shoots than on control shoots (81.3 vs. 90.9%), but survival was lower also on non-defoliated shoots where larvae had access only to current-year needles (78.8%). Data on larval feeding distribution showed a strong preference for mature needles. 5 Needle nitrogen concentration of current-year needles was 38% higher on defoliated trees than on non-defoliated trees. 6 It is concluded that defoliation affected larval performance primarily through the removal of the preferred type of needles and not because of an induced resistance. Effects of increased concentrations of allelochemicals in defoliated shoots, if present, were probably cancelled out by increased nitrogen concentrations.