Tree diversity enhances predation by birds but not by arthropods across climate gradients.

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.

Functional and phylogenetic relationships link predators to plant diversity via trophic and non-trophic pathways.

Human-induced biodiversity loss negatively affects ecosystem function, but the interactive effects of biodiversity change across trophic levels remain insufficiently understood. We sampled arboreal spiders and lepidopteran larvae across seasons in 2 years in a subtropical tree diversity experiment, and then disentangled the links between tree diversity and arthropod predator diversity by deconstructing the pathways among multiple components of diversity (taxonomic, phylogenetic and functional) with structural equation models. We found that herbivores were major mediators of plant species richness effects on abundance, species richness, functional and phylogenetic diversity of predators, while phylogenetic, functional and structural diversity of trees were also important mediators of this process. However, the strength and direction differed between functional, structural and phylogenetic diversity effects, indicating different underlying mechanisms for predator community assembly. Abundance and multiple diversity components of predators were consistently affected by tree functional diversity, indicating that the variation in structure and environment caused by plant functional composition might play key roles in predator community assembly. Our study highlights the importance of an integrated approach based on multiple biodiversity components in understanding the consequences of biodiversity loss in multitrophic communities.

Carbon-biodiversity relationships in a highly diverse subtropical forest.

Carbon-focused climate mitigation strategies are becoming increasingly important in forests. However, with ongoing biodiversity declines we require better knowledge of how much such strategies account for biodiversity. We particularly lack information across multiple trophic levels and on established forests, where the interplay between carbon stocks, stand age, and tree diversity might influence carbon-biodiversity relationships. Using a large dataset (>4600 heterotrophic species of 23 taxonomic groups) from secondary, subtropical forests, we tested how multitrophic diversity and diversity within trophic groups relate to aboveground, belowground, and total carbon stocks at different levels of tree species richness and stand age. Our study revealed that aboveground carbon, the key component of climate-based management, was largely unrelated to multitrophic diversity. By contrast, total carbon stocks-that is, including belowground carbon-emerged as a significant predictor of multitrophic diversity. Relationships were nonlinear and strongest for lower trophic levels, but nonsignificant for higher trophic level diversity. Tree species richness and stand age moderated these relationships, suggesting long-term regeneration of forests may be particularly effective in reconciling carbon and biodiversity targets. Our findings highlight that biodiversity benefits of climate-oriented management need to be evaluated carefully, and only maximizing aboveground carbon may fail to account for biodiversity conservation requirements.

Introduction of non-native Douglas fir reduces leaf damage on beech saplings and mature trees in European beech forests.

Recent ecological research suggests that, in general, mixtures are more resistant to insect herbivores and pathogens than monocultures. However, we know little about mixtures with non-native trees, where enemy release could lead to patterns that differ from commonly observed relationships among native species. This becomes particularly relevant when considering that adaptation strategies to climate change increasingly promote a larger share of non-native tree species, such as North American Douglas fir in Central Europe. We studied leaf damage on European beech (Fagus sylvatica) saplings and mature trees across a wide range of site conditions in monocultures and mixtures with phylogenetically distant conifers native Norway spruce (Picea abies) and non-native Douglas fir (Pseudotsuga menziesii). We analyzed leaf herbivory and pathogen damage in relation to tree diversity and composition effects, as well as effects of environmental factors and plant characteristics. We observed lower sapling herbivory and tree sucking damage on beech in non-native Douglas fir mixtures than in beech monocultures, probably due to a lower herbivore diversity on Douglas fir trees, and higher pathogen damage on beech saplings in Norway spruce than Douglas fir mixtures, possibly because of higher canopy openness. Our findings suggest that for low diversity gradients, tree diversity effects on leaf damage can strongly depend on tree species composition, in addition to modifications caused by feeding guild and tree ontogeny. Moreover, we found that nutrient capacity modulated the effects of tree diversity, composition, and environmental factors, with different responses in sites with low or high nutrient capacity. The existence of contrasting diversity effects based on tree species composition provides important information on our understanding of the relationships between tree diversity and plant-herbivore interactions in light of non-native tree species introductions. Especially with recent Norway spruce die-off, the planting of Douglas fir as replacement is likely to strongly increase in Central Europe. Our findings suggest that mixtures with Douglas fir could benefit the survival or growth rates of beech saplings and mature trees due to lower leaf damage, emphasizing the need to clearly identify and compare the potential benefits and ecological trade-offs of non-native tree species in forest management under ongoing environmental change.

Metabarcoding reveals that mixed forests mitigate negative effects of non-native trees on canopy arthropod diversity.

Averting climate change-induced forest diebacks increasingly relies on tree species planted outside of their natural range and on the addition of non-native tree species to mixed-species forests. However, the consequences of such changes for associated biodiversity remain poorly understood, especially for the forest canopy as a largely understudied forest stratum. Here, we used flight interception traps and a metabarcoding approach to study the taxonomic and functional (trophic guilds) composition and taxon richness of canopy arthropods. We sampled 15 monospecific and mixed stands of native European beech, native Norway spruce-planted outside its natural range-and non-native Douglas fir in northwest Germany. We found that the diversity of arthropods was lower in non-native Douglas fir compared with native beech stands. Taxon richness of herbivores was reduced by both conifer species. Other functional guilds, however, were not affected by stand type. Arthropod composition differed strongly between native broadleaved beech and monospecific coniferous (native spruce or non-native Douglas fir) stands, with less pronounced differences between the native and non-native conifers. Beech-conifer mixtures consistently hosted intermediate arthropod diversity and community composition compared with the respective monospecific stands. Moreover, arthropod diversity had a positive relationship with the number of canopy microhabitats. Our study shows that considering arthropod taxa of multiple functional groups reveals the multifaceted impact of non-native tree species on forest canopy arthropod communities. Contrasting with previous studies that primarily focused on the forest floor, we found that native beech hosts a rich diversity of arthropods, compared with lower diversity and distinct communities in economically attractive, and especially in non-native, conifers with few canopy microhabitats. Broadleaf-conifer mixtures did not perform better than native beech stands, but mitigated the negative effects of conifers, making such mixtures a compromise to foster both forest-associated diversity and economic yield.

Tree dissimilarity determines multi-dimensional beta-diversity of herbivores and carnivores via bottom-up effects.

Global biodiversity decline and its cascading effects through trophic interactions pose a severe threat to human society. Establishing the impacts of biodiversity decline requires a more thorough understanding of multi-trophic interactions and, more specifically, the effects that loss of diversity in primary producers has on multi-trophic community assembly. Within a synthetic conceptual framework for multi-trophic beta-diversity, we tested a series of hypotheses on neutral and niche-based bottom-up processes in assembling herbivore and carnivore communities in a subtropical forest using linear models, hieratical variance partitioning based on linear mixed-effects models (LMMs) and simulation. We found that the observed taxonomic, phylogenetic and functional beta-diversity of both herbivorous caterpillars and carnivorous spiders were significantly and positively related to tree dissimilarity. Linear models and variance partitioning for LMMs jointly suggested that as a result of bottom-up effects, producer dissimilarities were predominant in structuring consumer dissimilarity, the strength of which highly depended on the trophic dependencies on producers, the diversity facet examined, and data quality. Importantly, linear models for standardized beta-diversities against producer dissimilarities implied a transition between niche-based processes such as environmental filtering and competitive exclusion, which supports the role of bottom-up effect in determining consumer community assembly. These findings enrich our mechanistic understanding of the 'Diversity Begets Diversity' hypothesis and the complexity of higher-trophic community assembly, which is fundamental for sustainable biodiversity conservation and ecosystem management.

Tree species identity, canopy structure and prey availability differentially affect canopy spider diversity and trophic composition.

Forest canopies maintain a high proportion of arthropod diversity. The drivers that structure these communities, however, are poorly understood. Therefore, integrative research connecting tree species identity and environmental stand properties with taxonomic and functional community composition of canopy arthropods is required. In this study, we investigated how the taxonomic, functional and trophic composition of arboreal spider communities is affected by tree species composition and associated differences in canopy structure and prey availability in temperate forests. We sampled canopy spiders as well as their potential prey using insecticidal fogging in monospecific and mixed stands of native European beech, native Norway spruce and non-native Douglas fir. Trophic metrics were obtained from stable isotope analysis and structural canopy properties were assessed with mobile laser scanning. Monospecific native spruce stands promoted local canopy spider abundance and diversity, but native beech and beech-conifer mixtures had the highest diversity at landscape scale. Spider community composition differed between monospecific stands, with broadleaf-conifer mixtures mitigating these differences. Irrespective of tree species identity, spider abundance, taxonomic diversity, functional richness and isotopic richness increased in structurally heterogeneous canopies with high prey abundances, but functional evenness and trophic divergence decreased. Our study shows that canopy spiders are differentially affected by tree species identity, canopy structure and prey availability. Broadleaf-conifer mixtures mitigated negative effects of (non-native) conifers, but positive mixture effects were only evident at the landscape scale. Structurally heterogeneous canopies promoted the dominance of only specific trait clusters. This indicates that intermediate heterogeneity might result in high stability of ecological communities.

Tree communities and functional traits determine herbivore compositional turnover.

There are many factors known to drive species turnover, although the mechanisms by which these operate are less clear. Based on comprehensive datasets from the largest tree diversity experiment worldwide (BEF-China), we used shared herbivore species (zeta diversity) and multi-site generalized dissimilarity modelling to investigate the patterns and determinants of species turnover of Lepidoptera herbivores among study plots across a gradient in tree species richness. We found that zeta diversity declined sharply with an increasing number of study plots, with complete changes in caterpillar species composition observed even at the fine spatial scale of our study. Plant community characteristics rather than abiotic factors were found to play key roles in driving caterpillar compositional turnover, although these effects varied with an increasing number of study plots considered, due to the varying contributions of rare and common species to compositional turnover. Our study reveals details of the impact of phylogeny- and trait-mediated processes of trees on herbivore compositional turnover, which has implications for forest management and conservation and shows potential avenues for maintenance of heterogeneity in herbivore communities.

Host functional and phylogenetic composition rather than host diversity structure plant-herbivore networks.

Declining plant diversity alters ecological networks, such as plant-herbivore interactions. However, our knowledge of the potential mechanisms underlying effects of plant species loss on plant-herbivore network structure is still limited. We used DNA barcoding to identify herbivore-host plant associations along declining levels of tree diversity in a large-scale, subtropical biodiversity experiment. We tested for effects of tree species richness, host functional and phylogenetic diversity, and host functional (leaf trait) and phylogenetic composition on species, phylogenetic and network composition of herbivore communities. We found that phylogenetic host composition and related palatability/defence traits but not tree species richness significantly affected herbivore communities and interaction network complexity at both the species and community levels. Our study indicates that evolutionary dependencies and functional traits of host plants determine the composition of higher trophic levels and corresponding interaction networks in species-rich ecosystems. Our findings highlight that characteristics of the species lost have effects on ecosystem structure and functioning across trophic levels that cannot be predicted from mere reductions in species richness.

Tree species richness attenuates the positive relationship between mutualistic ant-hemipteran interactions and leaf chewer herbivory.

Interactions across trophic levels influence plant diversity effects on ecosystem functions, but the complexity of these interactions remains poorly explored. For example, the interplay between different interactions (e.g. mutualism, predation) might be an important moderator of biodiversity-ecosystem function relationships. We tested for relationships between trophobioses (facultative ant-hemipteran mutualism) and leaf chewer herbivory in a subtropical forest biodiversity experiment. We analysed trophobiosis and herbivory data of more than 10 000 trees along a tree species richness gradient. Against expectations, chewing damage was higher on trees with trophobioses. However, the net positive relationship between trophobioses and overall herbivory depended on tree species richness, being most pronounced at low richness. Our results point to indirect, positive effects of ant-tended sap suckers on leaf chewers, potentially by altering plant defences. Direct antagonistic relationships of trophobiotic ants and leaf-chewing herbivores-frequently reported to drive community-wide effects of trophobioses in other ecosystems-seemed less relevant. However, antagonistic interactions likely contributed to the attenuating effect of tree species richness, because trophobiotic ant and herbivore communities changed from monocultures to species-rich mixtures. Our findings, therefore, suggest that biodiversity loss might lead to complex changes in higher trophic level effects on ecosystem functions, mediated by both trophic and non-trophic interactions.

Tree diversity promotes generalist herbivore community patterns in a young subtropical forest experiment.

Stand diversification is considered a promising management approach to increasing the multifunctionality and ecological stability of forests. However, how tree diversity affects higher trophic levels and their role in regulating forest functioning is not well explored particularly for (sub)tropical regions. We analyzed the effects of tree species richness, community composition, and functional diversity on the abundance, species richness, and beta diversity of important functional groups of herbivores and predators in a large-scale forest biodiversity experiment in south-east China. Tree species richness promoted the abundance, but not the species richness, of the dominant, generalist herbivores (especially, adult leaf chewers), probably through diet mixing effects. In contrast, tree richness did not affect the abundance of more specialized herbivores (larval leaf chewers, sap suckers) or predators (web and hunting spiders), and only increased the species richness of larval chewers. Leaf chemical diversity was unrelated to the arthropod data, and leaf morphological diversity only positively affected oligophagous herbivore and hunting spider abundance. However, richness and abundance of all arthropods showed relationships with community-weighted leaf trait means (CWM). The effects of trait diversity and CWMs probably reflect specific nutritional or habitat requirements. This is supported by the strong effects of tree species composition and CWMs on herbivore and spider beta diversity. Although specialized herbivores are generally assumed to determine herbivore effects in species-rich forests, our study suggests that generalist herbivores can be crucial for trophic interactions. Our results indicate that promoting pest control through stand diversification might require a stronger focus on identifying the best-performing tree species mixtures.

Functional and phylogenetic diversity of woody plants drive herbivory in a highly diverse forest.

Biodiversity loss may alter ecosystem processes, such as herbivory, a key driver of ecological functions in species-rich (sub)tropical forests. However, the mechanisms underlying such biodiversity effects remain poorly explored, as mostly effects of species richness - a very basic biodiversity measure - have been studied. Here, we analyze to what extent the functional and phylogenetic diversity of woody plant communities affect herbivory along a diversity gradient in a subtropical forest. We assessed the relative effects of morphological and chemical leaf traits and of plant phylogenetic diversity on individual-level variation in herbivory of dominant woody plant species across 27 forest stands in south-east China. Individual-level variation in herbivory was best explained by multivariate, community-level diversity of leaf chemical traits, in combination with community-weighted means of single traits and species-specific phylodiversity measures. These findings deviate from those based solely on trait variation within individual species. Our results indicate a strong impact of generalist herbivores and highlight the need to assess food-web specialization to determine the direction of biodiversity effects. With increasing plant species loss, but particularly with the concomitant loss of functional and phylogenetic diversity in these forests, the impact of herbivores will probably decrease - with consequences for the herbivore-mediated regulation of ecosystem functions.

Tree diversity promotes functional dissimilarity and maintains functional richness despite species loss in predator assemblages.

The effects of species loss on ecosystems depend on the community's functional diversity (FD). However, how FD responds to environmental changes is poorly understood. This applies particularly to higher trophic levels, which regulate many ecosystem processes and are strongly affected by human-induced environmental changes. We analyzed how functional richness (FRic), evenness (FEve), and divergence (FDiv) of important generalist predators-epigeic spiders-are affected by changes in woody plant species richness, plant phylogenetic diversity, and stand age in highly diverse subtropical forests in China. FEve and FDiv of spiders increased with plant richness and stand age. FRic remained on a constant level despite decreasing spider species richness with increasing plant species richness. Plant phylogenetic diversity had no consistent effect on spider FD. The results contrast with the negative effect of diversity on spider species richness and suggest that functional redundancy among spiders decreased with increasing plant richness through non-random species loss. Moreover, increasing functional dissimilarity within spider assemblages with increasing plant richness indicates that the abundance distribution of predators in functional trait space affects ecological functions independent of predator species richness or the available trait space. While plant diversity is generally hypothesized to positively affect predators, our results only support this hypothesis for FD-and here particularly for trait distributions within the overall functional trait space-and not for patterns in species richness. Understanding the way predator assemblages affect ecosystem functions in such highly diverse, natural ecosystems thus requires explicit consideration of FD and its relationship with species richness.

Woody plant phylogenetic diversity mediates bottom-up control of arthropod biomass in species-rich forests.

Global change is predicted to cause non-random species loss in plant communities, with consequences for ecosystem functioning. However, beyond the simple effects of plant species richness, little is known about how plant diversity and its loss influence higher trophic levels, which are crucial to the functioning of many species-rich ecosystems. We analyzed to what extent woody plant phylogenetic diversity and species richness contribute to explaining the biomass and abundance of herbivorous and predatory arthropods in a species-rich forest in subtropical China. The biomass and abundance of leaf-chewing herbivores, and the biomass dispersion of herbivores within plots, increased with woody plant phylogenetic diversity. Woody plant species richness had much weaker effects on arthropods, but interacted with plant phylogenetic diversity to negatively affect the ratio of predator to herbivore biomass. Overall, our results point to a strong bottom-up control of functionally important herbivores mediated particularly by plant phylogenetic diversity, but do not support the general expectation that top-down predator effects increase with plant diversity. The observed effects appear to be driven primarily by increasing resource diversity rather than diversity-dependent primary productivity, as the latter did not affect arthropods. The strong effects of plant phylogenetic diversity and the overall weaker effects of plant species richness show that the diversity-dependence of ecosystem processes and interactions across trophic levels can depend fundamentally on non-random species associations. This has important implications for the regulation of ecosystem functions via trophic interaction pathways and for the way species loss may impact these pathways in species-rich forests.

Enhancing economic multifunctionality without compromising multidiversity and ecosystem multifunctionality via forest enrichment.

Enriching tree species-poor and less productive forests by introducing economically valuable species is a strategy proposed for achieving multipurpose forest management. However, empirical evidence from managed and mature forests on the impact of this enrichment on ecological (multidiversity and ecosystem multifunctionality) and economic dimensions remains scarce, particularly when nonnative species are used. Here, we propose and test a framework that integrates economic multifunctionality, encompassing timber production-oriented goals and resistance against disturbances, with multidiversity and ecosystem multifunctionality in European beech forest stands enriched with conifers. Our results show that enriched beech forest stands (~80 years old) can provide high levels of economic multifunctionality without compromising multidiversity and ecosystem multifunctionality. In comparison to pure beech stands, enriched stands with Douglas-fir supported win-win-win situations for these three dimensions. Our findings contribute to the discussion of integrating biodiversity, ecosystem, and economic functions, providing empirical evidence for future forest management.

Plant diversity enhances ecosystem multifunctionality via multitrophic diversity.

Ecosystem functioning depends on biodiversity at multiple trophic levels, yet relationships between multitrophic diversity and ecosystem multifunctionality have been poorly explored, with studies often focusing on individual trophic levels and functions and on specific ecosystem types. Here, we show that plant diversity can affect ecosystem functioning both directly and by affecting other trophic levels. Using data on 13 trophic groups and 13 ecosystem functions from two large biodiversity experiments-one representing temperate grasslands and the other subtropical forests-we found that plant diversity increases multifunctionality through elevated multitrophic diversity. Across both experiments, the association between multitrophic diversity and multifunctionality was stronger than the relationship between the diversity of individual trophic groups and multifunctionality. Our results also suggest that the role of multitrophic diversity is greater in forests than in grasslands. These findings imply that, to promote sustained ecosystem multifunctionality, conservation planning must consider the diversity of both plants and higher trophic levels.

Conifers and non-native tree species shift trophic niches of generalist arthropod predators in Central European beech forests.

BACKGROUND: Functional diversity is vital for forest ecosystem resilience in times of climate-induced forest diebacks. Admixing drought resistant non-native Douglas fir, as a partial replacement of climate-sensitive Norway spruce, to native beech forests in Europe appears promising for forest management, but possible consequences for associated biota and ecosystem functioning are poorly understood. To better link forest management and functional diversity of associated biota, we investigated the trophic niches (∆13C, ∆15N) of epigeic generalist predators (spiders and ground beetles) in mixed and pure stands of European beech, Norway spruce and non-native Douglas fir in north-west Germany. We assessed the multidimensional niche structure of arthropod predator communities using community-based isotopic metrics. RESULTS: Whilst arthropod ∆13C differed most between beech (high ∆13C) and coniferous stands (low ∆13C), ∆15N was lowest in non-native Douglas fir. Tree mixtures mitigated these effects. Further, conifers increased isotopic ranges and isotopic richness, which is linked to higher canopy openness and herb complexity. Isotopic divergence of ground beetles decreased with Douglas fir presence, and isotopic evenness of spiders in Douglas fir stands was lower in loamy sites with higher precipitation than in sandy, drier sites. CONCLUSIONS: We conclude that tree species and particularly non-native trees alter the trophic niche structure of generalist arthropod predators. Resource use and feeding niche breadth in non-native Douglas fir and native spruce differed significantly from native beech, with more decomposer-fueled and narrower feeding niches in beech stands (∆13C, isotopic ranges and richness). Arthropod predators in non-native Douglas fir, however, had shorter (∆15N) and simplified (isotopic divergence) food chains compared to native forest stands; especially under beneficial abiotic conditions (isotopic evenness). These findings indicate potential adverse effects of Douglas fir on functional diversity of generalist arthropod predators. As tree mixtures mitigated differences between beech and conifers, mixed stands including (non-native) conifers constitute a promising compromise between economic and conservational interests.

Multitrophic arthropod diversity mediates tree diversity effects on primary productivity.

Forests sustain 80% of terrestrial biodiversity and provide essential ecosystem services. Biodiversity experiments have demonstrated that plant diversity correlates with both primary productivity and higher trophic diversity. However, whether higher trophic diversity can mediate the effects of plant diversity on productivity remains unclear. Here, using 5 years of data on aboveground herbivorous, predatory and parasitoid arthropods along with tree growth data within a large-scale forest biodiversity experiment in southeast China, we provide evidence of multidirectional enhancement among the diversity of trees and higher trophic groups and tree productivity. We show that the effects of experimentally increased tree species richness were consistently positive for species richness and abundance of herbivores, predators and parasitoids. Richness effects decreased as trophic levels increased for species richness and abundance of all trophic groups. Multitrophic species richness and abundance of arthropods were important mediators of plant diversity effects on tree productivity, suggesting that optimizing forest management for increased carbon capture can be more effective when the diversity of higher trophic groups is promoted in concert with that of trees.

Plant traits affecting herbivory on tree recruits in highly diverse subtropical forests.

Differences in herbivory among woody species can greatly affect the functioning of forest ecosystems, particularly in species-rich (sub)tropical regions. However, the relative importance of the different plant traits which determine herbivore damage remains unclear. Defence traits can have strong effects on herbivory, but rarely studied geographical range characteristics could complement these effects through evolutionary associations with herbivores. Herein, we use a large number of morphological, chemical, phylogenetic and biogeographical characteristics to analyse interspecific differences in herbivory on tree saplings in subtropical China. Unexpectedly, we found no significant effects of chemical defence traits. Rather, herbivory was related to the plants' leaf morphology, local abundance and climatic niche characteristics, which together explained 70% of the interspecific variation in herbivory in phylogenetic regression. Our study indicates that besides defence traits and apparency to herbivores, previously neglected measures of large-scale geographical host distribution are important factors influencing local herbivory patterns among plant species.

Effects of enemy exclusion on biodiversity-productivity relationships in a subtropical forest experiment.

Interspecific niche complementarity is a key mechanism posited to explain positive species richness-productivity relationships in plant communities. However, the exact nature of the niche dimensions that plant species partition remains poorly known.Species may partition abiotic resources that limit their growth, but species may also be specialized with respect to their set of biotic interactions with other trophic levels, in particular with enemies including pathogens and consumers. The lower host densities present in more species-diverse plant communities may therefore result in smaller populations of specialized enemies, and in a smaller associated negative feedback these enemies exert on plant productivity.To test whether such host density-dependent effects of enemies drive diversity-productivity relationships in young forest stands, we experimentally manipulated leaf fungal pathogens and insect herbivores in a large subtropical forest biodiversity-ecosystem functioning experiment in China (BEF-China).We found that fungicide spraying of tree canopies removed the positive tree-species richness-productivity relationship present in untreated control plots. The tree species that contributed the most to this effect were the ones with the highest fungicide-induced growth increase in monoculture. Insecticide application did not cause comparable effects. Synthesis. Our findings suggest that tree species diversity may not only promote productivity by interspecific resource-niche partitioning but also by trophic niche partitioning. Most likely, partitioning occurred with respect to enemies such as pathogenic fungi. Alternatively, similar effects on tree growth would have occurred if fungicide had eliminated positive effects of a higher diversity of beneficial fungi (e.g. mycorrhizal symbionts) that may have occurred in mixed tree species communities.