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Recent long‐term tree biodiversity experiments have shown that diversity effects on productivity tend to strengthen over time, as complementarity among constituent species increases over the course of forest development. However, these community‐level metrics only account for the net outcome of multiple interactions among species and, thus, do not inform about the individual species' responses to diversity.In this study, using 11 years of growth records from a large diversity experiment, we explored how species respond to diversity based on their functional traits and those of their heterospecific neighbours over time and analysed their contribution to the community‐level overyielding.We show species‐specific responses to diversity, with fast‐growing deciduous species rapidly performing better in mixtures relative to monocultures, than slow‐growing evergreen species. Moreover, we find that species productivity in mixtures enhances over time as the proportion of slow‐growing evergreen species in the heterospecific neighbourhood increases. These patterns of response of species scale up and explain community overyielding, which occurs primarily in deciduous‐evergreen mixtures and is explained by the overyielding of deciduous species overcompensating the poor performance of evergreen species.This study sheds light on the temporal dynamics of species responses to diversity, which together help improve our understanding of community‐level overyielding over the course of stand development. Read the free Plain Language Summary for this article on the Journal blog. [ABSTRACT FROM AUTHOR]

There is increasing evidence that the strength of tree diversity effects on productivity varies considerably over the course of forest development. Evidence points to canopy closure and the subsequent self‐thinning as key phases of forest development during which positive diversity effects emerge. A number of studies have shown that self‐thinning can differ among species, and also in mixtures compared with monocultures. Yet, how diversity influences the process of canopy closure and self‐thinning remains poorly understood.In this study, using 11 years of growth and mortality records from a large diversity experiment, we fitted self‐thinning trajectories for 37 tree communities with equal initial densities and explored whether and how functional diversity and identity may affect these trajectories. We then examined whether the diversity effects on self‐thinning were influenced by differences in growth or in mortality.We found that tree communities' functional diversity and identity strongly influence the self‐thinning process. First, we observed that tree communities dominated by early successional species, and slow‐growing evergreens begin self‐thinning at a larger mean tree size. Second, we found that mixing species with contrasting resource‐use strategies, and the dominance of deciduous, fast‐growing species, reduce tree mortality rate in relation to mean tree size during self‐thinning (i.e. shallower self‐thinning slope). The lower rates of self‐thinning in these functionally diverse communities seem to be explained by both an increase in tree growth and a reduction in density‐related mortality simultaneously over time.Synthesis. Overall, this study highlights that increasing tree diversity has the potential to enhance forest productivity in the long term through a better performance during the self‐thinning process when competition for resources is most intense. [ABSTRACT FROM AUTHOR]

Although there is compelling evidence that tree diversity has an overall positive effect on forest productivity, there are important divergences among studies on the nature and strength of these diversity effects and their timing during forest stand development. To clarify conflicting results related to stand developmental stage, we explored how diversity effects on productivity change through time in a diversity experiment spanning 11 years. We show that the strength of diversity effects on productivity progressively increases through time, becoming significantly positive after 9 years. Moreover, we demonstrate that the strengthening of diversity effects is driven primarily by gradual increases in complementarity. We also show that mixing species with contrasting resource‐acquisition strategies, and the dominance of deciduous, fast‐developing species, promote positive diversity effects on productivity. Our results suggest that the canopy closure and subsequent stem exclusion phase are key for promoting niche complementarity in diverse tree communities. [ABSTRACT FROM AUTHOR]

In this ecoacoustic study we used the setting of a tropical tree diversity planted forest to analyze temporal patterns in the composition of soundscapes and to test the effects of tree species richness on associated biodiversity measured as acoustic diversity. The analysis of soundscapes offers easy, rapid and sustainable methods when assessing biodiversity. During the last years the quantification of regional or global acoustic variability in sounds and the analysis of different soundscapes has been evolving into an important tool for biodiversity conservation, especially since case studies confirmed a relationship between land-use management, forest structure and acoustic diversity. Here we analyzed soundscapes from two seasons (dry and rainy season) and aurally inspected a subset of audio recordings to describe temporal patterns in soundscape composition. Several acoustic indices were calculated and we performed a correlation analysis and a non-metric multidimensional scaling analysis to identify acoustic indices that: (i) were complementary to each other and such represented different aspects of the local soundscapes and (ii) related most strongly to differences in acoustic composition among tree species richness, season and day phase. Thus, we chose “High Frequency Cover,” “Bioacoustic Index,” and “Events Per Second” to test the hypothesis that acoustic diversity increases with increasing tree species richness. Monocultures differed significantly from polycultures during night recordings, with respect to High Frequency Cover. This index covers sounds above 8 kHz and thus represents part of the orthopteran community. We conclude that increasing tree species richness in a young tropical forest plantation had positive effects on the vocalizing communities. The strongest effects were found for acoustic activity of the orthopteran community. In contrast to birds, orthopterans have smaller home ranges, and are therefore important indicator species for small scale environmental conditions.

Abstract Soil C is the largest C pool in forest ecosystems that contributes to C sequestration and mitigates climate change. Tree diversity enhances forest productivity, so diversifying the tree species composition, notably in managed forests, could increase the quantity of organic matter being transferred to soils and alter other soil properties relevant to the C cycle. A ten‐year‐old tree diversity experiment was used to study the effects of tree identity and diversity (functional and taxonomic) on soils. Surface (0–10 cm) mineral soil was repeatedly measured for soil C concentration, C:N ratio, pH, moisture, and temperature in twenty‐four tree species mixtures and twelve corresponding monocultures (replicated in four blocks). Soil pH, moisture, and temperature responded to tree diversity and identity. Greater productivity in above‐ and below‐ground tree components did not increase soil C concentration. Soil pH increased and soil moisture decreased with functional diversity, more specifically, when species had different growth strategies and shade tolerances. Functional identity affected soil moisture and temperature, such that tree communities with more slow‐growing and shade‐tolerant species had greater soil moisture and temperature. Higher temperature was measured in communities with broadleaf‐deciduous species compared to communities with coniferous‐evergreen species. We conclude that long‐term soil C cycling in forest plantations will likely respond to changes in soil pH, moisture, and temperature that is mediated by tree species composition, since tree species affect these soil properties through their litter quality, water uptake, and physical control of soil microclimates.

Soil C is the largest C pool in forest ecosystems that contributes to C sequestration and mitigates climate change. Tree diversity enhances forest productivity, so diversifying the tree species composition, notably in managed forests, could increase the quantity of organic matter being transferred to soils and alter other soil properties relevant to the C cycle.A ten‐year‐old tree diversity experiment was used to study the effects of tree identity and diversity (functional and taxonomic) on soils. Surface (0–10 cm) mineral soil was repeatedly measured for soil C concentration, C:N ratio, pH, moisture, and temperature in twenty‐four tree species mixtures and twelve corresponding monocultures (replicated in four blocks).Soil pH, moisture, and temperature responded to tree diversity and identity. Greater productivity in above‐ and below‐ground tree components did not increase soil C concentration. Soil pH increased and soil moisture decreased with functional diversity, more specifically, when species had different growth strategies and shade tolerances. Functional identity affected soil moisture and temperature, such that tree communities with more slow‐growing and shade‐tolerant species had greater soil moisture and temperature. Higher temperature was measured in communities with broadleaf‐deciduous species compared to communities with coniferous‐evergreen species.We conclude that long‐term soil C cycling in forest plantations will likely respond to changes in soil pH, moisture, and temperature that is mediated by tree species composition, since tree species affect these soil properties through their litter quality, water uptake, and physical control of soil microclimates. [ABSTRACT FROM AUTHOR]

Global change exposes forest ecosystems to many risks including novel climatic conditions, increased frequency of climatic extremes and sudden emergence and spread of pests and pathogens. At the same time, forest landscape restoration has regained global attention as an integral strategy for climate change mitigation. Owing to unpredictable future risks and the need for new forests that provide multiple ecosystem services, mixed-species forests have been advocated for this purpose. However, the successful establishment of mixed forests requires intrinsic knowledge of biodiversity's role for forest ecosystem functioning. In this respect, a better understanding of tree-tree interactions and how they contribute to observed positive tree species richness effects on key ecosystem functions is critical. Here, we review the current knowledge of the underlying mechanisms of tree-tree interactions and argue that positive net biodiversity effects at the community scale may emerge from the dominance of positive over negative interactions at the local neighbourhood scale. In a second step, we demonstrate how tree-tree interactions and the immediate tree neighbourhood's role can be systematically assessed in a tree diversity experiment. The expected results will improve predictions about the effects of tree interactions on ecosystem functioning based on general principles. We argue that this knowledge is urgently required to guide the design of tree species mixtures for the successful establishment of newly planted forests. [ABSTRACT FROM AUTHOR]

Although both observational and experimental studies have shown that positive tree species diversity–productivity relationships are predominant in global forests, weak or the lack of tree species diversity and productivity relationships also exist. Growing evidence has revealed that ecosystem productivity depends more on the functional characteristics of species than on their number. However, exactly to what extent tree diversity effects on ecosystem productivity are influenced by the variability and composition of functional traits have rarely been tested both across and at given species richness (SR) levels.Here, we employed a meta‐analysis of global‐scale data from 59 tree diversity experiments to examine how the diversity and community‐weighted means (CWMs) of economics traits determine the outcomes of tree mixture effects on productivity across and within SR levels.We found that the positive effects of tree mixtures on productivity were strengthened by the increasing multidimensional functional dispersion (FDis) and the CWM of leaf nitrogen content both across and within two‐ and four‐species mixtures. Moreover, the multidimensional FDis and the CWM of leaf nitrogen content increased the complementarity effect rather than the selection effect.Synthesis. Our findings suggest that both diversity in the leaf economics spectrum and trait concentration on the 'fast' end of the spectrum strengthen biodiversity–ecosystem functioning relationships. This study provides mechanistic insights into the potent roles of plant economics traits, especially leaf nitrogen content, in determining the magnitude (and even directionality) of the biodiversity–ecosystem functioning relationships in forest ecosystems. [ABSTRACT FROM AUTHOR]

Background: Experimental manipulations of tree diversity have often found overyielding in mixed-species plantations. While most experiments are still in the early stages of stand development, the impacts of tree diversity are expected to accumulate over time. Here, I present findings from a 31-year-old tree diversity experiment (as of 2018) in Japan. Results: I find that the net diversity effect on stand biomass increased linearly through time. The species mixture achieved 64% greater biomass than the average monoculture biomass 31 years after planting. The complementarity effect was positive and increased exponentially with time. The selection effect was negative and decreased exponentially with time. In the early stages (≤ 3 years), the positive complementarity effect was explained by enhanced growths of early- and mid-successional species in the mixture. Later on (≥ 15 years), it was explained by their increased survival rates owing to vertical spatial partitioning — i.e. alleviation of self-thinning via canopy stratification. The negative selection effect resulted from suppressed growths of late-successional species in the bottom layer. Conclusions: The experiment provides pioneering evidence that the positive impacts of diversity-driven spatial partitioning on forest biomass can accumulate over multiple decades. The results indicate that forest biomass production and carbon sequestration can be enhanced by multispecies afforestation strategies. [ABSTRACT FROM AUTHOR]

While tree diversity has been proven to enhance above‐ground forest biomass in many instances, the effect on below‐ground biomass has often been neutral, or even negative. This raises the questions of whether above‐ground results are the product of a reduced allocation below‐ground in more diverse forests, which could imply a more efficient acquisition of below‐ground resources per unit root, and whether the effects of diversity have been correctly quantified and interpreted in the past.We addressed these issues using data from a tree‐based functional diversity experiment where 24 mixtures and their respective monocultures were analysed in a fully replicated design. We used species‐level measurements of above‐ and below‐ground biomass to assess how adding below‐ground biomass changes our interpretations regarding the roles of complementarity effects (niche partitioning or facilitation between species) and selection effects (dominance of high‐yielding species within mixtures).Mixtures showed overyielding of above‐ground biomass but not below‐ground. Although remaining significantly positive, this translated into diversity effects of a lesser magnitude when adding below‐ground biomass. However, species did not reduce their allocation to below‐ground biomass within mixtures compared to monocultures. Rather, this combination of above‐ground overyielding and below‐ground neutral effect was caused by the dominance in mixtures of species with a low below‐ground allocation strategy. Another consequence of these species dominance is that their actual contribution to total productivity within mixtures was overestimated when using only above‐ground biomass, due to their relatively small below‐ground biomass. This caused an overestimation of the selection or dominance effects, thereby causing an underestimation of the relative importance of complementarity effects.Synthesis. This study emphasizes the need to consider both above‐ and below‐ground biomass components when determining the origin and strength of diversity effects, or when attempting to link tree diversity to ecosystem services such as carbon sequestration. The statistical partitioning of net effects is a widely used method, but one which can lead to an overestimation of the role of individual dominant species when below‐ground biomass is not taken into account, hence diminishing the role of positive interactions between species. [ABSTRACT FROM AUTHOR]

Large‐scale conversion of tropical forests into oil palm monocultures has led to dramatic losses of biodiversity and ecosystem functions. While ecological restoration is urgently needed in many oil palm landscapes, there is a lack of scientific knowledge of sustainable management strategies.We established experimental tree islands of varying sizes (25 m2 to 1,600 m2) and diversity levels (1, 2, 3 and 6 species) in an oil palm plantation in Sumatra, Indonesia. Six native multi‐purpose tree species including Archidendron pauciflorum (Jengkol), Durio zibethinus (Durian), Parkia speciosa (Petai), Shorea leprosula (Meranti), Peronema canescens (Sungkai) and Dyera polyphylla We planted native trees in an oil palm landscape following a tree island pattern. The establishment success differed widely among the six planted species. The selection of particular species is a decisive factor to foster a positive relationship between diversity and tree growth. Planting larger tree islands (e.g. 1,600 square metres) is a better option to enhance tree growth, but tree‐palm competition implies a trade‐off between tree growth and palm oil production in the tree islands. [ABSTRACT FROM AUTHOR]

The widely observed positive relationship between plant diversity and ecosystem functioning is thought to be substantially driven by complementary resource use of plant species. Recent work suggests that biotic interactions among plants and between plants and soil organisms drive key aspects of resource use complementarity. Here, we provide a conceptual framework for integrating positive biotic interactions across guilds of organisms, more specifically between plants and mycorrhizal types, to explain resource use complementarity in plants and its consequences for plant competition. Our overarching hypothesis is that ecosystem functioning increases when more plant species associate with functionally dissimilar mycorrhizal fungi because differing mycorrhizal types will increase coverage of habitat space for and reduce competition among plants. We introduce a recently established field experiment (MyDiv) that uses different pools of tree species that associate with either arbuscular or ectomycorrhizal fungi to create orthogonal experimental gradients in tree species richness and mycorrhizal associations and present initial results. Finally, we discuss options for future mechanistic studies on resource use complementarity within MyDiv. We show how mycorrhizal types and biotic interactions in MyDiv can be used in the future to test novel questions regarding the mechanisms underlying biodiversity–ecosystem function relationships. [ABSTRACT FROM AUTHOR]

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 'BEF-China,' 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. [ABSTRACT FROM AUTHOR]