Differential Responses of Herbivores and Herbivory to Management in Temperate European Beech

Forest management not only affects biodiversity but also might alter ecosystem processes mediated by the organisms, i.e. herbivory the removal of plant biomass by plant-eating insects and other arthropod groups. Aiming at revealing general relationships between forest management and herbivory we investigated aboveground arthropod herbivory in 105 plots dominated by European beech in three different regions in Germany in the sun-exposed canopy of mature beech trees and on beech saplings in the understorey. We separately assessed damage by different guilds of herbivores, i.e. chewing, sucking and scraping herbivores, gall-forming insects and mites, and leaf-mining insects. We asked whether herbivory differs among different forest management regimes (unmanaged, uneven-aged managed, even-aged managed) and among age-classes within even-aged forests. We further tested for consistency of relationships between regions, strata and herbivore guilds. On average, almost 80% of beech leaves showed herbivory damage, and about 6% of leaf area was consumed. Chewing damage was most common, whereas leaf sucking and scraping damage were very rare. Damage was generally greater in the canopy than in the understorey, in particular for chewing and scraping damage, and the occurrence of mines. There was little difference in herbivory among differently managed forests and the effects of management on damage differed among regions, strata and damage types. Covariates such as wood volume, tree density and plant diversity weakly influenced herbivory, and effects differed between herbivory types. We conclude that despite of the relatively low number of species attacking beech; arthropod herbivory on beech is generally high. We further conclude that responses of herbivory to forest management are multifaceted and environmental factors such as forest structure variables affecting in particular microclimatic conditions are more likely to explain the variability in herbivory among beech forest plots.     

Tree diversity drives diversity of arthropod herbivores,but successional stage mediates detritivores

The high tree diversity of subtropical forests is linked to the biodiversity of other trophic levels. Disentangling the effects of tree species richness and composition, forest age, and stand structure on higher trophic levels in a forest landscape is important for understanding the factors that promote biodiversity and ecosystem functioning. Using a plot network spanning gradients of tree diversity and secondary succession in subtropical forest, we tested the effects of tree community characteristics (species richness and composition) and forest succession (stand age) on arthropod community characteristics (morphotype diversity, abundance and composition) of four arthropod functional groups. We posit that these gradients differentially affect the arthropod functional groups, which mediates the diversity, composition, and abundance of arthropods in subtropical forests. We found that herbivore richness was positively related to tree species richness. Furthermore, the composition of herbivore communities was associated with tree species composition. In contrast, detritivore richness and composition was associated with stand age instead of tree diversity. Predator and pollinator richness and abundance were not strongly related to either gradient, although positive trends with tree species richness were found for predators. The weaker effect of tree diversity on predators suggests a cascading diversity effect from trees to herbivores to predators. Our results suggest that arthropod diversity in a subtropical forest reflects the net outcome of complex interactions among variables associated with tree diversity and stand age. Despite this complexity, there are clear linkages between the overall richness and composition of tree and arthropod communities, in particular herbivores, demonstrating that these trophic levels directly impact each other.     

Herbivory on planted oak seedlings across a habitat edge created by timber harvest

Edge habitats create environmental gradients that affect plant community composition and herbivore behavior. Silvicultural disturbance creates edge habitat with direct (via changes in light) and indirect (via changes in herbivore behavior) consequences for the growth and survival of tree seedlings, and thus, the composition of the future forest stands. Herbivores, particularly ungulates, can be a major limiting factor in oak regeneration, and silvicultural disturbance may alter the abundance or behavior of herbivores following harvest. We measured the severity of herbivory on experimentally planted white (Quercus alba) and black oak (Quercus velutina) seedlings by white-tailed deer (Odocoileus virginianus) and eastern cottontail rabbits (Sylvilagus floridanus), as well as foliar damage from insects, across gradients created by clearcuts in a deciduous forest in Indiana, USA. Overall browse pressure on oaks was low in our study. Nonetheless, spatial variation in herbivory depended on herbivore taxa; herbivory by rabbits was highest inside harvest openings, whereas foliar damage by insects peaked in the forest. Intensity of deer herbivory was constant across the edge. In addition, we observed indirect interactions among herbivore species mediated by a seedling’s browsing history. Herbivore damage by deer was positively related to past browsing by rabbits, and foliar damage from insects was positively related to past browsing by both deer and rabbits. Increasing woody plant competition reduced herbivory on seedlings by both deer and rabbits. Given the lack of spatial variability in deer herbivory and low overall herbivory by rabbits, we suspect that interactions between timber harvesting and herbivory did not have a strong impact on oak seedlings at our study sites.     

Comparisons of arthropod assemblages on an invasive and native trees: abundance,diversity and damage

The success of exotic plants may be due to lower herbivore loads than those on native plants (Enemies Release Hypothesis). Predictions of this hypothesis include lower herbivore abundances, diversity, and damage on introduced plant species compared to native ones. Greater density or diversity of predators and parasitoids on exotic versus native plants may also reduce regulation of exotic plants by herbivores. To test these predictions, we measured arthropod abundance, arthropod diversity, and foliar damage on invasive Chinese tallow tree (Triadica sebifera) and three native tree species: silver maple (Acer saccharinum), sycamore (Platanus occidentalis), and sweetgum (Liquidambar styraciflua). Arthropod samples were collected with canopy sweep nets from six 20 year old monoculture plots of each species at a southeast Texas site. A total of 2,700 individuals and 285 species of arthropods were caught. Overall, the species richness and abundance of arthropods on tallow tree were similar to the natives. But, ordination (NMS) showed community composition differed on tallow tree compared to all three native trees. It supported an arthropod community that had relatively lower herbivore abundance but relatively more predator species compared to the native species examined. Leaves were collected to determine damage. Tallow tree experienced less mining damage than native trees. The results of this study supported the Enemies Release Hypothesis predictions that tallow tree would have low herbivore loads which may contribute to its invasive success. Moreover, a shift in the arthropod community to fewer herbivores without a reduction in predators may further limit regulation of this exotic species by herbivores in its introduced range.     

Tree species diversity influences herbivore abundance and damage: meta-analysis of long-term forest experiments

Plant monocultures are commonly believed to be more susceptible to herbivore attacks than stands composed of several plant species. However, few studies have experimentally tested the effects of tree species diversity on herbivory. In this paper, we present a meta-analysis of uniformly collected data on insect herbivore abundance and damage on three tree species (silver birch, black alder and sessile oak) from seven long-term forest diversity experiments in boreal and temperate forest zones. Our aim was to compare the effects of forest diversity on herbivores belonging to different feeding guilds and inhabiting different tree species. At the same time we also examined the variation in herbivore responses due to tree age and sampling period within the season, the effects of experimental design (plot size and planting density) and the stability of herbivore responses over time. Herbivore responses varied significantly both among insect feeding guilds and among host tree species. Among insect feeding guilds, only leaf miner densities were consistently lower and less variable in mixed stands as compared to tree monocultures regardless of the host tree species. The responses of other herbivores to forest diversity depended largely on host tree species. Insect herbivory on birch was significantly lower in mixtures than in birch monocultures, whereas insect herbivory on oak and alder was higher in mixtures than in oak and alder monocultures. The effects of tree species diversity were also more pronounced in older trees, in the earlier part of the season, at larger plots and at lower planting density. Overall our results demonstrate that forest diversity does not generally and uniformly reduce insect herbivory and suggest instead that insect herbivore responses to forest diversity are highly variable and strongly dependent on the host tree species and other stand characteristics as well as on the type of the herbivore.     

Species and structural diversity of church forests in a fragmented Ethiopian Highland landscape

Question: Thousands of small isolated forest fragments remain around churches (“church forests”) in the almost completely deforested Ethiopian Highlands. We questioned how the forest structure and composition varied with altitude, forest area and human influence. Location: South Gondar, Amhara National Regional State, Northern Ethiopia. Methods: The structure and species composition was assessed for 810 plots in 28 church forests. All woody plants were inventoried, identified and measured (stem diameter) in seven to 56 10 m x 10‐m plots per forest. Results: In total, 168 woody species were recorded, of which 160 were indigeneous. The basal area decreased with tree harvest intensity; understorey and middle‐storey density (<5 cm DBH trees) decreased with grazing; overstorey density (>5 cm DBH trees) increased with altitude. The dominance of a small set of species increased with altitude and grazing intensity. Species richness decreased with altitude, mainly due to variation in the richness of the overstorey community. Moreover, species richness in the understorey decreased with grazing intensity. Conclusions: We show how tree harvesting intensity, grazing intensity and altitude contribute to observed variations in forest structure, composition and species richness. Species richness was, however, not related to forest area. Our study emphasizes the significant role played by the remaining church forests for conservation of woody plant species in North Ethiopian Highlands, and the need to protect these forests for plant species conservation purposes.     

Delayed local responses of downy birch to damage by leafminers and leafrollers

Recent findings suggest that impacts of endemic herbivory on forest ecosystems over the long term may exceed impacts of herbivore outbreaks. However, responses of trees to minor and local damage imposed by small arthropod herbivores, especially by those mining or skeletonising individual leaves, remain poorly understood. We studied the delayed effects of injuries by several leafmining and leafrolling insects on the performance of downy birch shoots. Insect feeding did not affect survival of shoots or survival of individual axillary buds in long shoots. In the year following the damage, shoots produced an average of 13.8% more biomass than undamaged shoots of the same tree. The magnitude of this effect increased with an increase in the leaf area injured during the previous year, but it did not differ among four localities in subarctic and boreo‐nemoral forests, between herbivore feeding guilds, or among herbivores imposing damage in early, mid and late summer. We also found that herbivores attacked the next‐year foliage produced by damaged shoots less frequently than they attacked the next‐year foliage produced by undamaged shoots of the same tree. Thus, our study demonstrated delayed local compensatory growth and increased antiherbivore defence in downy birch shoots following local damage by insect feeding. We suggest that this pattern reflects evolutionary adaptations of plants to permanently acting minor, dispersed and spatially unpredictable damage imposed by endemic herbivory. Local responses are less costly and represent a more sustainable strategy to maintain plant fitness under low levels of herbivory than constitutive resistance or systemic responses.     

Insect attraction to herbivore-induced beech volatiles under different forest management regimes

Insect herbivore enemies such as parasitoids and predators are important in controlling herbivore pests. From agricultural systems we know that land-use intensification can negatively impact biological control as an important ecosystem service. The aim of our study was to investigate the importance of management regime for natural enemy pressure and biological control possibilities in forests dominated by European beech. We hypothesize that the volatile blend released from herbivore-infested beech trees functions as a signal, attracting parasitoids and herbivore enemies. Furthermore, we hypothesize that forest management regime influences the composition of species attracted by these herbivore-induced beech volatiles. We installed flight-interception traps next to Lymantria dispar caterpillar-infested young beech trees releasing herbivore-induced volatiles and next to non-infested control trees. Significantly more parasitoids were captured next to caterpillar-infested trees compared to non-infested controls, irrespective of forest type. However, the composition of the trophic guilds in the traps did vary in response to forest management regime. While the proportion of chewing insects was highest in non-managed forests, the proportion of sucking insects peaked in forests with low management and of parasitoids in young, highly managed, forest stands. Neither the number of naturally occurring beech saplings nor herbivory levels in the proximity of our experiment affected the abundance and diversity of parasitoids caught. Our data show that herbivore-induced beech volatiles attract herbivore enemies under field conditions. They further suggest that differences in the structural complexity of forests as a consequence of management regime only play a minor role in parasitoid activity and thus in indirect tree defense.     

Sapling herbivory,invertebrate herbivores and predators across a natural tree diversity gradient in Germany’s largest connected deciduous forest

Tree species-rich forests are hypothesised to be less susceptible to insect herbivores, but so far herbivory–diversity relationships have rarely been tested for tree saplings, and no such study has been published for deciduous forests in Central Europe. We expected that diverse tree communities reduce the probability of detection of host plants and increase abundance of predators, thereby reducing herbivory. We examined levels of herbivory suffered by beech (Fagus sylvatica L.) and maple saplings (Acer pseudoplatanus L. and Acer platanoides L.) across a tree species diversity gradient within Germany’s largest remaining deciduous forest area, and investigated whether simple beech or mixed stands were less prone to damage caused by herbivorous insects. Leaf area loss and the frequency of galls and mines were recorded for 1,040 saplings (>13,000 leaves) in June and August 2006. In addition, relative abundance of predators was assessed to test for potential top-down control. Leaf area loss was generally higher in the two species of maple compared to beech saplings, while only beech showed a decline in damage caused by leaf-chewing herbivores across the tree diversity gradient. No significant patterns were found for galls and mines. Relative abundance of predators on beech showed a seasonal response and increased on species-rich plots in June, suggesting higher biological control. We conclude that, in temperate deciduous forests, herbivory–tree diversity relationships are significant, but are tree species-dependent with bottom-up and top-down control as possible mechanisms. In contrast to maple, beech profits from growing in a neighbourhood of higher tree richness, which implies that species identity effects may be of greater importance than tree diversity effects per se. Hence, herbivory on beech appeared to be mediated bottom-up by resource concentration in the sampled forest stands, as well as regulated top-down through biocontrol by natural enemies. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorised users.     

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 tree hollow characteristics and forest structure on saproxylic beetle diversity in tree hollows in managed forests in a regional comparison

Tree hollows are among the rarest habitats in today''s Central European managed forests but are considered key structures for high biodiversity in forests. To analyze and compare the effects of tree hollow characteristics and forest structure on diversity of saproxylic beetles in tree hollows in differently structured managed forests, we examined between 41 and 50 tree hollows in beech trees in each of three state forest management districts in Germany. During the two‐year study, we collected 283 saproxylic beetle species (5880 individuals; 22% threatened species), using emergence traps. At small spatial scales, the size of hollow entrance and the number of surrounding microhabitat structures positively influenced beetle diversity, while the stage of wood mould decomposition had a negative influence, across all three forest districts. We utilized forest inventory data to analyze the effects of forest structure in radii of 50–500 m around tree hollows on saproxylic beetle diversity in the hollows. At these larger spatial scales, the three forest management districts differed remarkably regarding the parameters that influenced saproxylic beetle diversity in tree hollows. In Ebrach, characterized by mostly deciduous trees, the amount of dead wood positively influenced beetle diversity. In the mostly coniferous Fichtelberg forest district, with highly isolated tree hollows, in contrast, only the proportion of beech trees around the focal tree hollows showed a positive influence on beetle diversity. In Kelheim, characterized by mixed forest stands, there were no significant relationships between forest structure and beetle diversity in tree hollows. In this study, the same local tree hollow parameters influenced saproxylic beetle diversity in all three study regions, while parameters of forest structure at larger spatial scales differed in their importance, depending on tree‐species composition.     

Global patterns of insect herbivory in gap and understorey environments,and their implications for woody plant carbon storage

Insect herbivory is thought to favour carbon allocation to storage in juveniles of shade‐tolerant trees. This argument assumes that insect herbivory in the understorey is sufficiently intense as to select for storage; however, understoreys might be less attractive to insect herbivores than canopy gaps, because of low resource availability and – at temperate latitudes – low temperatures. Although empirical studies show that shade‐tolerant species in tropical forests do allocate more photosynthate to storage than their light‐demanding associates, the same pattern has not been consistently observed in temperate forests. Does this reflect a latitudinal trend in the relative activity of insect herbivory in gap versus understorey environments? To date there has been no global review of the effect of light environment on insect herbivory in forests. We postulated that if temperature is the primary factor limiting insect herbivory, the effect of gaps on rates of insect herbivory should be more evident in temperate than in tropical forests; due to low growing season temperatures in the oceanic temperate forests of the Southern Hemisphere, the effect of gaps on insect herbivory rates should in turn be stronger there than in the more continental temperate climates of the Northern Hemisphere. We examined global patterns of insect herbivory in gaps versus understories through meta‐analysis of 87 conspecific comparisons of leaf damage in contrasting light environments. Overall, insect herbivory in gaps was significantly higher than in the understorey; insect herbivory was 50% higher in gaps than in understoreys of tropical forests but did not differ significantly between gaps and understories in temperate forests of either hemisphere. Results are consistent with the idea that low resource availability – and not temperature – limits insect herbivore activity in forest understoreys, especially in the tropics, and suggest the selective influence of insect herbivory on late‐successional tree species may have been over‐estimated.     

Effective distance of volatile cues for plant–plant communication in beech

In response to volatiles emitted from a plant infested by herbivorous arthropods, neighboring undamaged conspecific plants become better defended against herbivores; this is referred to as plant‒plant communication. Although plant‒plant communication occurs in a wide range of plant species, most studies have focused on herbaceous plants. Here, we investigated plant‒plant communication in beech trees in two experimental plantations in 2018 and one plantation in 2019. Approximately 20% of the leaves of a beech tree were clipped in half in the spring seasons of 2018 and 2019 (clipped tree). The damage levels to leaves in the surrounding undamaged beech trees were evaluated 90 days after the clipping (assay trees). In both years, the damage levels decreased with a reduction in the distance from the clipped tree. In 2019, we also recorded the damage levels of trees that were not exposed to volatiles (nonexposed trees) as control trees and found that those that were located <5 m away from clipped trees had significantly less leaf damage than nonexposed trees. By using a gas chromatograph–mass spectrometer, ten and eight volatile compounds were detected in the headspaces of clipped and unclipped leaves, respectively. Among them, the amount of (Z)‐3‐hexenyl acetate in clipped leaves was significantly higher than that in nonclipped leaves. Our result suggests that green leaf volatiles such as (Z)‐3‐hexenol and (Z)‐3‐hexenyl acetate and other volatile organic compounds emitted from clipped trees induced defenses in the neighboring trees within the 5 m radius. The effective distances of plant‒plant communication in trees were discussed from the viewpoint of the arthropod community structure in forest ecosystems.     

Tree diversity drives associational resistance to herbivory at both forest edge and interior

Tree diversity is increasingly acknowledged as an important driver of insect herbivory. However, there is still a debate about the direction of associational effects that can range from associational resistance (i.e., less damage in mixed stands than in monocultures) to the opposite, associational susceptibility. Discrepancies among published studies may be due to the overlooked effect of spatially dependent processes such as tree location within forests. We addressed this issue by measuring crown defoliation and leaf damage made by different guilds of insect herbivores on oaks growing among conspecific versus heterospecific neighbors at forest edges versus interior, in two closed sites in SW France forests. Overall, oaks were significantly less defoliated among heterospecific neighbors (i.e., associational resistance), at both forest edge and interior. At the leaf level, guild diversity and leaf miner herbivory significantly increased with tree diversity regardless of oak location within stands. Other guilds showed no clear response to tree diversity or oak location. We showed that herbivore response to tree diversity varied among insect feeding guilds but not between forest edges and interior, with inconsistent patterns between sites. Importantly, we show that oaks were more defoliated in pure oak plots than in mixed plots at both edge and forest interior and that, on average, defoliation decreased with increasing tree diversity from one to seven species. We conclude that edge conditions could be interacting with tree diversity to regulate insect defoliation, but future investigations are needed to integrate them into the management of temperate forests, notably by better understanding the role of the landscape context.     

Additive and non‐additive effects of birch genotypic diversity on arthropod herbivory in a long‐term field experiment

Herbivores are important drivers of plant population dynamics and community composition in natural and managed systems. Intraspecific genetic diversity of long‐lived plants like trees might shape patterns of herbivory by different guilds of herbivores that trees experience through time. However, previous studies on plant genetic diversity effects on herbivores have been largely short‐term. We investigated how tree genotypic variation and diversity influence herbivory of silver birch Betula pendula in a long‐term field experiment. Using clones of eight genotypes, we constructed experimental plots consisting of one, two, four or eight genotypes, and measured damage by five guilds of arthropod herbivores twice a year over three different years (four, six and nine years after the experiment was established). Genotypes varied significantly for most types of herbivore damage, but genotype resistance rankings often shifted over time, and none of the clones was more resistant than all others to all types of herbivores. At the plot level, birch genotypic diversity had significant positive additive effect on leaf rollers and negative non‐additive effects on chewing herbivores and gall makers. In contrast, leaf‐mining and leaf‐tying damage was not influenced by birch genotypic diversity. Within diverse plots, the direction of genotypic diversity effects varied depending on birch genotype, some having lower and some having higher herbivory in mixed stands. This research highlights the importance of long‐term studies including different feeding guilds of herbivores to understand the effects of plant genetic diversity on arthropod communities. Different responses of various feeding guilds to genotypic diversity and shifts in resistance of individual genotypes over time indicate that genotypic mixtures are unlikely to result in overall reduction in herbivory over time.     

Does insect herbivory on oak depend on the diversity of tree stands?

Studies on the effects of plant diversity on insect herbivory have produced conflicting results. Plant diversity has been reported to cause positive and negative responses of herbivores. Explanations for these conflicting responses include not only various population-level processes but also changes in plant quality that lead to changes in herbivore performance. In a tree diversity experiment, we investigated the effects of tree diversity on insect herbivory on oak in general and whether the effects of tree diversity on herbivore damage are reflected by the performance (leaf consumption, growth) of the generalist herbivore Lymantria dispar. Our study showed that the feeding damage caused by naturally occurring herbivores on oak trees decreased with increasing diversity of tree stands. The performance of L. dispar on oak leaves was not affected by tree diversity, neither in field nor laboratory experiments. Our results can be explained by the various processes behind the hypothesis of associational resistance.     

Leaf damage and functional traits along a successional gradient in Brazilian tropical dry forests

Herbivory has significant impacts on individual plants and plant communities, both at ecological and evolutionary time scales. In this context, this study aims to evaluate herbivore damage and its relationship with leaf chemical and structural traits, nutritional status, and forest structural complexity along a successional gradient. We predicted that trees in early successional stages support conservative traits related to drought tolerance (high specific leaf mass and phenolics), whereas trees in light-limited, late successional stages tend to enhance light acquisition strategies (high nitrogen content). We sampled 261 trees from 26 species in 15 plots (50 × 20 m; five per successional stage). From each tree, twenty leaves were collected for leaf trait measures. Phenolic content increased whereas specific leaf mass and nitrogen content decreased from early to late stages. However, leaf damage did not differ among successional stages. Our results partially corroborate the hypothesis that early successional plants in tropical dry forests exhibit leaf traits involved in the conservative use of water. The unexpected decrease in nitrogen content along the chronosequence is likely related to the fact that thinner leaves with low specific leaf mass could have less nitrogen-containing mesophyll per unit area. Mechanisms affecting herbivory intensity varied across scales: at the species level, leaf damage was negatively correlated with tannin concentration and specific leaf mass; at the plot level, leaf damage was positively affected by forest structural complexity. Herbivory patterns in tropical forests are difficult to detect because abiotic factors and multiple top-down and bottom-up forces directly and indirectly affect herbivores.     

Seedling Mortality and Herbivory Damage in Subtropical and Temperate Populations: Testing the Hypothesis of Higher Herbivore Pressure Toward the Tropics

Herbivory rates are generally thought to be higher in tropical than in temperate forests. Nevertheless, tests of this biogeographic prediction by comparing a single plant species across a tropical-temperate range are scarce. Here, we compare herbivore damage between subtropical and temperate populations of the evergreen tree Aextoxicon punctatum (Olivillo), distributed between 30° and 43° S along the Pacific margin of Chile. To assess the impact of herbivory on Olivillo seedlings, we set up 29 experimental plots, 1.5 × 3 m: 16 in forests of Fray Jorge National Park (subtropical latitude), and 13 in Guabún, Chiloé Island (temperate latitude). Half of each plot was fenced around with chicken wire, to exclude small mammals, and the other half was left unfenced. In each half of the plots we planted 16 seedlings of Olivillo in December 2003, with a total of 928 plants. Seedling survival, leaf production and herbivory by invertebrates were monitored over the next 16 mo. Small mammal herbivores killed ca 30 percent of seedlings in both sites. Nevertheless, invertebrate herbivory was greater in the temperate forest, thus contradicting the expected trend of increasing herbivore impact toward the tropics. Seedling growth was greater in subtropical forest suggesting better conditions for tree growth or that higher invertebrate herbivory depressed seedling growth in the temperate forest. Invertebrate herbivory increased toward temperate latitudes while small mammal herbivory was similar in both sites. We suggest that comparison of single species can be useful to test generalizations about latitudinal patterns and allow disentangling factors controlling herbivory patterns across communities.     

Beech forest management does not affect the infestation rate of the beech scale Cryptococcus fagisuga across three regions in Germany

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Contrasting effects of tree diversity on young tree growth and resistance to insect herbivores across three biodiversity experiments

Tree diversity is an important driver of forest ecosystem functioning, hypothesised to enhance tree growth and resistance to herbivores. To test this, we assessed the relative importance of tree species richness and functional diversity on tree height growth and insect herbivore damage across three tree diversity experiments in Finland, France and Germany, established within the last fifteen years. These experiments encompass species richness gradients from monocultures up to five species mixtures, with compositions drawn from a pool of eleven tree species. Tree height growth and total insect herbivory were evaluated at both the tree species and forest plot scales. Trees in mixtures tended to grow taller, but on average received more insect herbivory relative to monocultures. Gradients of tree species richness or functional diversity had only weak impact on the magnitude of these effects. Community weighted means of specific leaf area alone captured diversity effects on tree height growth, with stronger positive effects of diversity in mixtures with high community SLA. Tree species‐specific responses were highly variable. No species significantly benefited both in terms of increased growth and reduced herbivory when grown in mixtures. More species showed positive height growth responses in mixed assemblages, but only the two exotic conifers experienced associational resistance to herbivores. This large‐scale study shows that tree height growth in young forest plantations tends to be higher in species mixtures than in monocultures, but incremental increases in functional diversity have, at best, weak marginal growth benefits. Moreover, there appear to be contrasting effects at forest plot versus individual species scales. Thus, while some species show lower herbivore damage in mixtures, this is not a consistent trend and contrasts the higher overall damage in mixtures observed at the forest plot scale. To improve both tree growth and resistance to herbivores in tree species mixtures seems therefore challenging.