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Recent droughts have strongly impacted forest ecosystems and are projected to increase in frequency, intensity, and duration in the future together with continued warming. While evidence suggests that tree diversity can regulate drought impacts in natural forests, few studies examine whether mixed tree plantations are more resistant to the impacts of severe droughts. Using natural variations in leaf carbon (C) and nitrogen (N) isotopic ratios, that is δ 13 C and δ 15 N, as proxies for drought response, we analyzed the effects of tree species richness on the functional responses of tree plantations to the pan-European 2018 summer drought in seven European tree diversity experiments. We found that leaf δ 13 C decreased with increasing tree species richness, indicating less drought stress. This effect was not related to drought intensity, nor desiccation tolerance of the tree species. Leaf δ 15 N increased with drought intensity, indicating a shift toward more open N cycling as water availability diminishes. Additionally, drought intensity was observed to alter the influence of tree species richness on leaf δ 15 N from weakly negative under low drought intensity to weakly positive under high drought intensity. Overall, our findings suggest that dual leaf isotope analysis helps understand the interaction between drought, nutrients, and species richness., (© 2024 The Authors. New Phytologist © 2024 New Phytologist Foundation.)

Tree diversity can promote both predator abundance and diversity. However, whether this translates into increased predation and top-down control of herbivores across predator taxonomic groups and contrasting environmental conditions remains unresolved. We used a global network of tree diversity experiments (TreeDivNet) spread across three continents and three biomes to test the effects of tree species richness on predation across varying climatic conditions of temperature and precipitation. We recorded bird and arthropod predation attempts on plasticine caterpillars in monocultures and tree species mixtures. Both tree species richness and temperature increased predation by birds but not by arthropods. Furthermore, the effects of tree species richness on predation were consistent across the studied climatic gradient. Our findings provide evidence that tree diversity strengthens top-down control of insect herbivores by birds, underscoring the need to implement conservation strategies that safeguard tree diversity to sustain ecosystem services provided by natural enemies in forests., (© 2024 The Authors. Ecology Letters published by John Wiley & Sons Ltd.)

Many plant species produce multiple leaf flushes during the growing season, which might have major consequences for within-plant variation in chemistry and species interactions. Yet, we lack a theoretical or empirical framework for how differences among leaf flushes might shape variation in damage by insects and diseases. We assessed the impact of leaf flush identity on leaf chemistry, insect attack and pathogen infection on the pedunculate oak Quercus robur by sampling leaves from each leaf flush in 20 populations across seven European countries during an entire growing season. The first leaf flush had higher levels of primary compounds, and lower levels of secondary compounds, than the second flush, whereas plant chemistry was highly variable in the third flush. Insect attack decreased from the first to the third flush, whereas infection by oak powdery mildew was lowest on leaves from the first flush. The relationship between plant chemistry, insect attack and pathogen infection varied strongly among leaf flushes and seasons. Our findings demonstrate the importance of considering differences among leaf flushes for our understanding of within-tree variation in chemistry, insect attack and disease levels, something particularly relevant given the expected increase in the number of leaf flushes with climate change., (© 2022 The Authors. New Phytologist © 2022 New Phytologist Foundation.)

Forest pests can cause massive ecological and economic damage worldwide. Ecologically sound solutions to diminish forest insect pest impacts include the use of their natural enemies, such as predators and parasitoids, as well as entomopathogenic fungi, bacteria or viruses. Phytochemical compounds mediate most interactions between these organisms, but knowledge of such chemically mediated multitrophic relationships is still at its infancy for forest systems, particularly when compared to agricultural systems. Here, we highlight the main gaps in how phytochemicals of forest trees facilitate or interfere with trophic interactions between trees, insect herbivores, and interacting organisms including predators, parasitoids and microbes. We propose future avenues of research on phytochemical-based biocontrol of forest pests taking into account the characteristics of trees and forests., (Copyright © 2021 The Authors. Published by Elsevier Inc. All rights reserved.)

Ecological research conducted over the past five decades has shown that increasing tree species richness at forest stands can improve tree resistance to insect pest damage. However, the commonality of this finding is still under debate. In this review, we provide a quantitative assessment (i.e., a meta-analysis) of tree diversity effects on insect herbivory and discuss plausible mechanisms underlying the observed patterns. We provide recommendations and working hypotheses that can serve to lay the groundwork for research to come. Based on more than 600 study cases, our quantitative review indicates that insect herbivory was, on average, lower in mixed forest stands than in pure stands, but these diversity effects were contingent on herbivore diet breadth and tree species composition. In particular, tree species diversity mainly reduced damage of specialist insect herbivores in mixed stands with phylogenetically distant tree species. Overall, our findings provide essential guidance for forest pest management.

Both species and genetic diversity of plant communities can affect insect herbivores, but a few studies have compared the effects of both diversity levels within the same experimental context. We compared the effects of tree species and genetic diversity on abundance, species richness, and β-diversity of leaf-miner communities associated with silver birch using two long-term forest diversity experiments in Finland where birch trees were planted in monocultures and mixtures of birch genotypes or other trees species. Although both abundance and species richness of leaf miners differed among birch genotypes at the tree level, birch genetic diversity had no significant effect on miner abundance and species richness at the plot level. Instead, birch genetic diversity affected leaf-miner β-diversity with species turnover being higher among trees within genotypic mixtures than among trees within monoclonal plots. In contrast, tree species diversity had a significant negative effect on both leaf-miner abundance and species richness at plot level, but no effect on miner β-diversity. Significant tree species diversity effects on leaf-miner abundance and species richness were found only in plots with high tree density. We have demonstrated that plant species and genetic diversity play important but contrasting roles in structuring associated herbivore communities. Tree species diversity largely affects miner abundance and species richness, whereas tree genetic diversity affects miner β-diversity. These results have important implications for conservation and management of woodlands.

Forests with higher tree diversity are often assumed to be more resistant to insect herbivores but whether this effect depends on climatic conditions is so far poorly understood. In particular, a forest's resistance to herbivory may depend on mean annual temperature (MAT) as a key driver of plant and insect phenology. We carried out a global meta-analysis on regression coefficients between tree diversity and four aspects of insect herbivory, namely herbivore damage, abundance, incidence rate and species richness. To test for a potential shift of tree diversity effects along a global gradient of MAT we applied mixed-effects models and estimated grand mean effect sizes and the influence of MAT, experimental vs. observational studies and herbivores diet breadth. There was no overall effect of tree diversity on the pooled effect sizes of insect herbivore damage, abundance and incidence rate. However, when analysed separately, we found positive grand mean effect sizes for herbivore abundance and species richness. For herbivore damage and incidence rate we found a significant but opposing shift along a gradient of MAT indicating that with increasing MAT diversity effects on herbivore damage tend towards associational resistance whereas diversity effects on incidence rates tend towards associational susceptibility. Our results contradict previous meta-analyses reporting overall associational resistance to insect herbivores in mixed forests. Instead, we report that tree diversity effects on insect herbivores can follow a biogeographic pattern calling for further in-depth studies in this field., Competing Interests: The authors have declared that no competing interests exist.

Forest pest damage is expected to increase with global change. Tree diversity could mitigate this impact, but unambiguous demonstration of the diversity-resistance relationship is lacking in semi-natural mature forests. We used a network of 208 forest plots sampled along two orthogonal gradients of increasing tree species richness and latitudes to assess total tree defoliation in Europe. We found a positive relationship between tree species richness and resistance to insect herbivores: overall damage to broadleaved species significantly decreased with the number of tree species in mature forests. This pattern of associational resistance was frequently observed across tree species and countries, irrespective of their climate. These findings confirm the greater potential of mixed forests to face future biotic disturbances in a changing world., (© 2016 The Authors.)

The impact of invasive herbivore species may be lower in more diverse plant communities due to mechanisms of associational resistance. According to the "resource concentration hypothesis" the amount and accessibility of host plants is reduced in diverse plant communities, thus limiting the exploitation of resources by consumers. In addition, the "natural enemy hypothesis" suggests that richer plant assemblages provide natural enemies with more complementary resources and habitats, thus promoting top down regulation of herbivores. We tested these two hypotheses by comparing crown damage by the invasive Asian chestnut gall wasp (Dryocosmus kuriphilus) on chestnut trees (Castanea sativa) in pure and mixed stands in Italy. We estimated the defoliation on 70 chestnut trees in 15 mature stands sampled in the same region along a gradient of tree species richness ranging from one species (chestnut monocultures) to four species (mixtures of chestnut and three broadleaved species). Chestnut defoliation was significantly lower in stands with higher tree diversity. Damage on individual chestnut trees decreased with increasing height of neighboring, heterospecific trees. These results suggest that conservation biological control method based on tree species mixtures might help to reduce the impact of the Asian chestnut gall.