Biodiversity experiments have shown that species loss reduces ecosystem functioning in grassland. To test whether this result can be extrapolated to forests, the main contributors to terrestrial primary productivity, requires large-scale experiments. We manipulated tree species richness by planting more than 150,000 trees in plots with 1 to 16 species. Simulating multiple extinction scenarios, we found that richness strongly increased stand-level productivity. After 8 years, 16-species mixtures had accumulated over twice the amount of carbon found in average monocultures and similar amounts as those of two commercial monocultures. Species richness effects were strongly associated with functional and phylogenetic diversity. A shrub addition treatment reduced tree productivity, but this reduction was smaller at high shrub species richness. Our results encourage multispecies afforestation strategies to restore biodiversity and mitigate climate change.

Summary1. Biodiversity-ecosystem functioning (BEF) experiments address ecosystem-level consequences of species loss by comparing communities of high species richness with communities from which species have been gradually eliminated. BEF experiments originally started with microcosms in the laboratory and with grassland ecosystems. A new frontier in experimental BEF research is manipulating tree diversity in forest ecosystems, compelling researchers to think big and comprehensively. 2. We present and discuss some of the major issues to be considered in the design of BEF experiments with trees and illustrate these with a new forest biodiversity experiment established in subtropical China (Xingangshan, Jiangxi Province) in 2009/2010. Using a pool of 40 tree species, extinction scenarios were simulated with tree richness levels of 1, 2, 4, 8 and 16 species on a total of 566 plots of 25Á8 9 25Á8 m each. 3. The goal of this experiment is to estimate effects of tree and shrub species richness on carbon storage and soil erosion; therefore, the experiment was established on sloped terrain. The following important design choices were made: (i) establishing many small rather than fewer larger plots, (ii) using high planting density and random mixing of species rather than lower planting density and patchwise mixing of species, (iii) establishing a map of the initial 'ecoscape' to characterize site heterogeneity before the onset of biodiversity effects and (iv) manipulating tree species richness not only in random but also in trait-oriented extinction scenarios. 4. Data management and analysis are particularly challenging in BEF experiments with their hierarchical designs nesting individuals within-species populations within plots within-species compositions. Statistical analysis best proceeds by partitioning these random terms into fixed-term contrasts, for example, species composition into contrasts for species richness and the presence of particular functional groups, which can then be tested against the remaining random variation among compositions. 5. We conclude that forest BEF experiments provide exciting and timely research options. They especially require careful thinking to allow multiple disciplines to measure and analyse data jointly and effectively. Achiev- This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited. 2014, 5, 74-89 doi: 10.1111/2041-210X.12126 ing specific research goals and synergy with previous experiments involves trade-offs between different designs and requires manifold design decisions. Methods in Ecology andEvolution

(2011). Community assembly during secondary forest succession in a Chinese subtropical forest. Ecological Monographs, 81(1): 25-41. Community assembly during secondary forest succession in a Chinese subtropical forest Abstract Subtropical broad-leaved forests in southeastern China support a high diversity of woody plants. Using a comparative study design with 30330 m plots (n=27) from five successional stages (<20, <40, <60, <80, and ≤80 yr), we investigated how the gradient in species composition reflects underlying processes of community assembly. In particular, we tested whether species richness of adult trees and shrubs decreased or increased and assessed to which degree this pattern was caused by negative density dependence or continuous immigration over time. Furthermore, we tested whether rare species were increasingly enriched and the species composition of adult trees and shrubs became more similar to species composition of seedlings during the course of succession. We counted the individuals of all adult species and shrubs >1 m in height in each plot and counted all woody recruits (bank of all seedlings ≤1 m in height) in each central 10310 m quadrant of each plot. In addition, we measured a number of environmental variables (elevation, slope, aspect, soil moisture, pH, C, N, and C/N ratio) and biotic structural variables (height and cover of layers). Adult species richness varied from 25 to 69 species per plot, and in total 148 woody species from 46 families were recorded. There was a clear successional gradient in species composition as revealed by nonmetric multidimensional scaling (NMDS), but only a poor differentiation of different successional stages with respect to particular species. Adult richness per 100 individuals (rarefaction method) increased with successional stage. None of the measured abiotic variables were significantly correlated with adult species richness. We found no evidence that rare species were responsible for the increasing adult species richness, as richness of rare species among both adults and recruits was independent of the successional stage. Furthermore, the similarity between established adults and recruits did not increase with successional stage. There was a constant number of recruit species and also of exclusive recruit species, i.e., those that had not been present as adult individuals, across all successional stages, suggesting a continuous random immigration over time. variables were significantly correlated with adult species richness. We found no evidence that 41 rare species were responsible for the increasing adult species richness, as richness of rare 42 species amongst both adults and recruits was independent of the successional stage.

Local‐scale microclimatic conditions in forest understoreys play a key role in shaping the composition, diversity and function of these ecosystems. Consequently, understanding what drives variation in forest microclimate is critical to forecasting ecosystem responses to global change, particularly in the tropics where many species already operate close to their thermal limits and rapid land‐use transformation is profoundly altering local environments. Yet our ability to characterize forest microclimate at ecologically meaningful scales remains limited, as understorey conditions cannot be directly measured from outside the canopy. To address this challenge, we established a network of microclimate sensors across a land‐use intensity gradient spanning from old‐growth forests to oil‐palm plantations in Borneo. We then combined these observations with high‐resolution airborne laser scanning data to characterize how topography and canopy structure shape variation in microclimate both locally and across the landscape. In the processes, we generated high‐resolution microclimate surfaces spanning over 350 km2, which we used to explore the potential impacts of habitat degradation on forest regeneration under both current and future climate scenarios. We found that topography and vegetation structure were strong predictors of local microclimate, with elevation and terrain curvature primarily constraining daily mean temperatures and vapour pressure deficit (VPD), whereas canopy height had a clear dampening effect on microclimate extremes. This buffering effect was particularly pronounced on wind‐exposed slopes but tended to saturate once canopy height exceeded 20 m—suggesting that despite intensive logging, secondary forests remain largely thermally buffered. Nonetheless, at a landscape‐scale microclimate was highly heterogeneous, with maximum daily temperatures ranging between 24.2 and 37.2°C and VPD spanning two orders of magnitude. Based on this, we estimate that by the end of the century forest regeneration could be hampered in degraded secondary forests that characterize much of Borneo's lowlands if temperatures continue to rise following projected trends.

Tropical forests play a major role in the carbon cycle of the terrestrial biosphere. Recent field studies have provided detailed descriptions of the carbon cycle of mature tropical forests, but logged or secondary forests have received much less attention. Here, we report the first measures of total net primary productivity (NPP) and its allocation along a disturbance gradient from old-growth forests to moderately and heavily logged forests in Malaysian Borneo. We measured the main NPP components (woody, fine root and canopy NPP) in old-growth (n = 6) and logged (n = 5) 1 ha forest plots. Overall, the total NPP did not differ between old-growth and logged forest (13.5 ± 0.5 and 15.7 ± 1.5 Mg C ha year respectively). However, logged forests allocated significantly higher fraction into woody NPP at the expense of the canopy NPP (42% and 48% into woody and canopy NPP, respectively, in old-growth forest vs 66% and 23% in logged forest). When controlling for local stand structure, NPP in logged forest stands was 41% higher, and woody NPP was 150% higher than in old-growth stands with similar basal area, but this was offset by structure effects (higher gap frequency and absence of large trees in logged forest). This pattern was not driven by species turnover: the average woody NPP of all species groups within logged forest (pioneers, nonpioneers, species unique to logged plots and species shared with old-growth plots) was similar. Hence, below a threshold of very heavy disturbance, logged forests can exhibit higher NPP and higher allocation to wood; such shifts in carbon cycling persist for decades after the logging event. Given that the majority of tropical forest biome has experienced some degree of logging, our results demonstrate that logging can cause substantial shifts in carbon production and allocation in tropical forests.

Experimental forest plantations to study biodiversity-ecosystem functioning (BEF) relationships have recently been established in different regions of the world, but subtropical biomes have not been covered so far. Here, we report about the initial survivorship of 26 tree species in the first such experiment in subtropical China. In the context of the joint Sino-German-Swiss Research Unit "BEFChina," 271 experimental forest plots were established using 24 naturally occurring tree species and two native commercial conifers. Based on the survival inventories carried out in November 2009 and June 2010, the overall survival rate was 87 % after the first 14 months. Generalized mixed-effects models showed that survival rates of seedlings were significantly affected by species richness, the species' leaf habit (deciduous or evergreen), species identity, planting date, and altitude. In the first survey, seedling establishment success decreased with increasing richness levels, a tendency that disappeared in the second survey after replanting. Though evergreen species performed less well than deciduous species with establishment rates of 84 versus 93 % in the second survey, their planting success exceeded the general expectation for subtropical broad-leaved evergreen species. These results have important implications for establishing mixed-species plantations for diversity conservation and improvement of ecosystem functioning in the Chinese subtropics and elsewhere. Additional costs associated with mixed-species plantations as compared to conventional plantations also demonstrate the potential of upscaling BEF experiments to large-scale afforestation projects. June 2010, the overall survival rate was 87% after the first 14 months. Generalized mixed-effects 22 models showed that survival rates of seedlings were significantly affected by species richness, the 23 species' leaf habit (deciduous or evergreen), species identity, planting date and altitude. In the first 24 survey, seedling establishment success decreased with increasing richness levels, a tendency that 25 disappeared in the second survey after replanting. Deciduous species had a significantly higher survival 26 than evergreen species (93% vs. 84% in the second survey). These results have implications for 27 establishing Though evergreen species performed less well than deciduous species with establishment 28 rates of 84% vs. 93% in the second survey, their planting success exceeded the general expectation for 29 subtropical broad-leave evergreen species. These results have important implications for establishing 30 mixed species plantations for diversity conservation and improvement of ecosystem functioning in the 31Chinese subtropics and elsewhere. Additional costs associated with mixed -species as compared to 32 conventional plantations also demonstrates the potential of upscaling BEF experiments to large-scale 33 afforestation projects. 34 35

Summary Plant functional traits regulate ecosystem functions but little is known about how co‐occurring gradients of land use and edaphic conditions influence their expression. We test how gradients of logging disturbance and soil properties relate to community‐weighted mean traits in logged and old‐growth tropical forests in Borneo. We studied 32 physical, chemical and physiological traits from 284 tree species in eight 1 ha plots and measured long‐term soil nutrient supplies and plant‐available nutrients. Logged plots had greater values for traits that drive carbon capture and growth, whilst old‐growth forests had greater values for structural and persistence traits. Although disturbance was the primary driver of trait expression, soil nutrients explained a statistically independent axis of variation linked to leaf size and nutrient concentration. Soil characteristics influenced trait expression via nutrient availability, nutrient pools, and pH. Our finding, that traits have dissimilar responses to land use and soil resource availability, provides robust evidence for the need to consider the abiotic context of logging when predicting plant functional diversity across human‐modified tropical forests. The detection of two independent axes was facilitated by the measurement of many more functional traits than have been examined in previous studies.

Predatory arthropods can exert strong top-down control on ecosystem functions. However, despite extensive theory and experimental manipulations of predator diversity, our knowledge about relationships between plant and predator diversity—and thus information on the relevance of experimental findings—for species-rich, natural ecosystems is limited. We studied activity abundance and species richness of epigeic spiders in a highly diverse forest ecosystem in subtropical China across 27 forest stands which formed a gradient in tree diversity of 25–69 species per plot. The enemies hypothesis predicts higher predator abundance and diversity, and concomitantly more effective top-down control of food webs, with increasing plant diversity. However, in our study, activity abundance and observed species richness of spiders decreased with increasing tree species richness. There was only a weak, non-significant relationship with tree richness when spider richness was rarefied, i.e. corrected for different total abundances of spiders. Only foraging guild richness (i.e. the diversity of hunting modes) of spiders was positively related to tree species richness. Plant species richness in the herb layer had no significant effects on spiders. Our results thus provide little support for the enemies hypothesis—derived from studies in less diverse ecosystems—of a positive relationship between predator and plant diversity. Our findings for an important group of generalist predators question whether stronger top-down control of food webs can be expected in the more plant diverse stands of our forest ecosystem. Biotic interactions could play important roles in mediating the observed relationships between spider and plant diversity, but further testing is required for a more detailed mechanistic understanding. Our findings have implications for evaluating the way in which theoretical predictions and experimental findings of functional predator effects apply to species-rich forest ecosystems, in which trophic interactions are often considered to be of crucial importance for the maintenance of high plant diversity.