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Abstract Forest canopy complexity (i.e., the three‐dimensional structure of the canopy) is often associated with increased species diversity as well as high primary productivity across natural forests. However, canopy complexity, tree diversity, and productivity are often confounded in natural forests, and the mechanisms of these relationships remain unclear. Here, we used two large tree diversity experiments in North America to assess three hypotheses: (1) increasing tree diversity leads to increased canopy complexity, (2) canopy complexity is positively related to tree productivity, and (3) the relationship between tree diversity and tree productivity is indirect and driven by the positive effects of canopy complexity. We found that increasing tree diversity from monocultures to mixtures of 12 species increases canopy complexity and productivity by up to 71% and 73%, respectively. Moreover, structural equation modeling indicates that the effects of tree diversity on productivity are indirect and mediated primarily by changes in internal canopy complexity. Ultimately, we suggest that increasing canopy complexity can be a major mechanism by which tree diversity enhances productivity in young forests. 

Abstract Structural diversity—the volume and physical arrangement of vegetation within the three‐dimensional (3D) space of ecosystems—is a predictor of ecosystem function that can be measured at large scales with remote sensing. However, the landscape composition and configuration of structural diversity across macrosystems have not been well described. Using a relatively recently developed method to quantify landscape composition and configuration of continuous habitat or terrain, we propose the application of gradient surface metrics (GSMs) to quantify landscape patterns of structural diversity and provide insights into how its spatial pattern relates to ecosystem function. We first applied an example set of GSMs that represent landscape heterogeneity, dominance, and edge density to Lidar‐derived structural diversity within 28 forested landscapes at National Ecological Observatory Network (NEON) sites. Second, we tested for forest type, geographic location, and climate drivers of macroscale variation in GSMs of structural diversity (GSM‐SD). Third, we demonstrated the utility of these metrics for understanding spatial patterns of ecosystem function in a case study with NDVI, a proxy of productivity. We found that GSM‐SD varied in landscapes within macrosystems, with forest type, geographic location, and climate being significantly related to some but not all metrics. We also found that dominance of high peaks of height and vertical complexity of canopy vegetation and the heterogeneity of the vertical complexity and coefficient of variation of canopy vegetation height within 120‐m patches were negatively correlated with NDVI across the 28 NEON sites. However, forest type always had a significant interaction term between these GSM‐SD and NDVI relationships. Our study demonstrates that GSMs are useful to describe the landscape composition and configuration of structural diversity and its relationship with productivity that warrants further consideration for spatially motivated management decisions. 

Abstract Plant diversity effects on community productivity often increase over time. Whether the strengthening of diversity effects is caused by temporal shifts in species-level overyielding (i.e., higher species-level productivity in diverse communities compared with monocultures) remains unclear. Here, using data from 65 grassland and forest biodiversity experiments, we show that the temporal strength of diversity effects at the community scale is underpinned by temporal changes in the species that yield. These temporal trends of species-level overyielding are shaped by plant ecological strategies, which can be quantitatively delimited by functional traits. In grasslands, the temporal strengthening of biodiversity effects on community productivity was associated with increasing biomass overyielding of resource-conservative species increasing over time, and with overyielding of species characterized by fast resource acquisition either decreasing or increasing. In forests, temporal trends in species overyielding differ when considering above- versus belowground resource acquisition strategies. Overyielding in stem growth decreased for species with high light capture capacity but increased for those with high soil resource acquisition capacity. Our results imply that a diversity of species with different, and potentially complementary, ecological strategies is beneficial for maintaining community productivity over time in both grassland and forest ecosystems. 

Summary Decades of studies have demonstrated links between biodiversity and ecosystem functioning, yet the generality of the relationships and the underlying mechanisms remain unclear, especially for forest ecosystems.Using 11 tree‐diversity experiments, we tested tree species richness–community productivity relationships and the role of arbuscular (AM) or ectomycorrhizal (ECM) fungal‐associated tree species in these relationships.Tree species richness had a positive effect on community productivity across experiments, modified by the diversity of tree mycorrhizal associations. In communities with both AM and ECM trees, species richness showed positive effects on community productivity, which could have resulted from complementarity between AM and ECM trees. Moreover, both AM and ECM trees were more productive in mixed communities with both AM and ECM trees than in communities assembled by their own mycorrhizal type of trees. In communities containing only ECM trees, species richness had a significant positive effect on productivity, whereas species richness did not show any significant effects on productivity in communities containing only AM trees.Our study provides novel explanations for variations in diversity–productivity relationships by suggesting that tree–mycorrhiza interactions can shape productivity in mixed‐species forest ecosystems. 

Abstract One mechanism proposed to explain high species diversity in tropical systems is strong negative conspecific density dependence (CDD), which reduces recruitment of juveniles in proximity to conspecific adult plants. Although evidence shows that plant-specific soil pathogens can drive negative CDD, trees also form key mutualisms with mycorrhizal fungi, which may counteract these effects. Across 43 large-scale forest plots worldwide, we tested whether ectomycorrhizal tree species exhibit weaker negative CDD than arbuscular mycorrhizal tree species. We further tested for conmycorrhizal density dependence (CMDD) to test for benefit from shared mutualists. We found that the strength of CDD varies systematically with mycorrhizal type, with ectomycorrhizal tree species exhibiting higher sapling densities with increasing adult densities than arbuscular mycorrhizal tree species. Moreover, we found evidence of positive CMDD for tree species of both mycorrhizal types. Collectively, these findings indicate that mycorrhizal interactions likely play a foundational role in global forest diversity patterns and structure. 

Abstract 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. 

Abstract Human actions are decreasing the diversity and complexity of forests, and a mechanistic understanding of how these changes affect predators is needed to maintain ecosystem services, including pest regulation. Using a large‐scale tree diversity experiment, we investigate how spiders respond to trees growing in plots of single or mixed species combinations (4 or 12) by repeatedly sampling 540 trees spanning 15 species. In 2019 (6 years post‐establishment), spider responses to tree diversity varied by tree species. By 2021, diversity had a more consistently positive effect, with trees in 4‐ or 12‐species plots supporting 23% or 50% more spiders, respectively, compared to conspecifics in monocultures. Spiders showed stronger tree species preferences in late summer, and the positive impact of plot diversity doubled. In early summer, the positive diversity effect was tied to higher canopy cover in diverse plots, leading to higher spider densities. This indirect path strengthened in late summer, with an additional direct effect of plot diversity on spiders. Prey availability was higher in diverse plots but was not tied to spider density. Overall, diverse plots supported more predators, partly by increasing available habitat. Adopting planting strategies focused on species mixtures may better maintain higher trophic levels and ecosystem functions. 

Abstract Efforts to catalog global biodiversity have often focused on aboveground taxonomic diversity, with limited consideration of belowground communities. However, diversity aboveground may influence the diversity of belowground communities and vice versa. In addition to taxonomic diversity, the structural diversity of plant communities may be related to the diversity of soil bacterial and fungal communities, which drive important ecosystem processes but are difficult to characterize across broad spatial scales. In forests, canopy structural diversity may influence soil microorganisms through its effects on ecosystem productivity and root architecture, and via associations between canopy structure, stand age, and species richness. Given that structural diversity is one of the few types of diversity that can be readily measured remotely (e.g., using light detection and ranging—LiDAR), establishing links between structural and microbial diversity could facilitate the detection of belowground biodiversity hotspots. We investigated the potential for using remotely sensed information about forest structural diversity as a predictor of soil microbial community richness and composition. We calculated LiDAR‐derived metrics of structural diversity as well as a suite of stand and soil properties from 38 forested plots across the central hardwoods region of Indiana, USA, to test whether forest canopy structure is linked with the community richness and diversity of four key soil microbial groups: bacteria, fungi, arbuscular mycorrhizal (AM) fungi, and ectomycorrhizal (EM) fungi. We found that the density of canopy vegetation is positively associated with the taxonomic richness (alpha diversity) of EM fungi, independent of changes in plant taxonomic richness. Further, structural diversity metrics were significantly correlated with the overall community composition of bacteria, EM, and total fungal communities. However, soil properties were the strongest predictors of variation in the taxonomic richness and community composition of microbial communities in comparison with structural diversity and tree species diversity. As remote sensing tools and algorithms are rapidly advancing, these results may have important implications for the use of remote sensing of vegetation structural diversity for management and restoration practices aimed at preserving belowground biodiversity. 

Abstract Identifying the primary controls of particulate (POM) and mineral‐associated organic matter (MAOM) content in soils is critical for determining future stocks of soil carbon (C) and nitrogen (N) across the globe. However, drivers of these soil organic matter fractions are likely to vary among ecosystems in response to climate, soil type and the composition of local biological communities.We tested how soil factors, climate and plant–fungal associations influenced the distribution and concentrations of C and N in MAOM and POM in seven temperate forests in the National Ecological Observatory Network (NEON) across the eastern United States. Samples of upper mineral horizon soil within each forest were collected in plots representing a gradient of dominant tree–mycorrhizal association, allowing us to test how plant and microbial communities influenced POM and MAOM across sites differing in climate and soil conditions.We found that concentrations of C and N in soil organic matter were primarily driven by soil mineralogy, but the relative abundance of MAOM versus POM C was strongly linked to plot‐level mycorrhizal dominance. Furthermore, the effect of dominant tree mycorrhizal type on the distribution of N among POM and MAOM fractions was sensitive to local climate: in cooler sites, an increasing proportion of ectomycorrhizal‐associated trees was associated with lower proportions of N in MAOM, but in warmer sites, we found the reverse. As an indicator of soil carbon age, we measured radiocarbon in the MAOM fraction but found that within and across sites, Δ¹⁴C was unrelated to mycorrhizal dominance, climate, or soil factors, suggesting that additional site‐specific factors may be primary determinants of long‐term SOM persistence.Synthesis. Our results indicate that while soil mineralogy primarily controls SOM C and N concentrations, the distribution of SOM among density fractions depends on the composition of vegetation and microbial communities, with these effects varying across sites with distinct climates. We also suggest that within biomes, the age of mineral‐associated soil carbon is not clearly linked to the factors that control concentrations of MAOM C and N. 

Summary First principles predict that diversity at one trophic level often begets diversity at other levels, suggesting plant and mycorrhizal fungal diversity should be coupled. Local‐scale studies have shown positive coupling between the two, but the association is less consistent when extended to larger spatial and temporal scales. These inconsistencies are likely due to divergent relationships of different mycorrhizal fungal guilds to plant diversity, scale dependency, and a lack of coordinated sampling efforts. Given that mycorrhizal fungi play a central role in plant productivity and nutrient cycling, as well as ecosystem responses to global change, an improved understanding of the coupling between plant and mycorrhizal fungal diversity across scales will reduce uncertainties in predicting the ecosystem consequences of species gains and losses.