Through complementarity interactions, mixedspecies forests can be more productive than monocultures, and this effect can increase with tree-species richness. However, this is not always the case. This review examines the processes and stand structural attributes that can influence diversity-productivity relationships (DPRs); how they influence resource availability, resource uptake, and resource-use efficiency; and also describes some important differences between tree-diversity versus grassland-diversity experiments. The size of the complementarity effects caused by these processes and stand structures varies along spatial and temporal gradients in resource availability and climate. These spatial and temporal dynamics have now been examined in many studies, and the general patterns are summarized using a simple framework; complementarity is predicted to increase as the availability of resource BX^declines (or climatic condition X becomes harsher) if the species interactions improve the availability, uptake, or use efficiency of resource X (or interactions improve climatic condition X). Importantly, this framework differs from the stress-gradient hypothesis to account for a wider range of inter-specific plant interactions (not only facilitation) by considering contrasting methods used to quantify species interactions while accounting for stand structure. In addition, complementarity (as opposed to facilitation) for a given species combination can increase as growing conditions improve in forests, contrary to predictions of the stressgradient hypothesis with regards to facilitation. This review indicates that while the effects of tree-species diversity on growth and other forest functions are now receiving a lot of attention, far less is known about the effects of structural diversity on growth or forest functioning. Direct measurements of the processes, as opposed to focusing mainly on growth responses, could greatly contribute to our understanding of structural diversity effects.

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Mixed-species plantations of Eucalyptus with a nitrogen (N 2 ) fixing species have the potential to increase productivity while maintaining soil fertility, compared to Eucalyptus monocultures. However, it is difficult to predict combinations of species and sites that will lead to these benefits. We review the processes and interactions occurring in mixed plantations, and the influence of species or site attributes, to aid the selection of successful combinations of species and sites. Successful mixtures, where productivity is increased over that of monocultures, have often developed stratified canopies, such that the less shade-tolerant species overtops the more shade-tolerant species. Successful mixtures also have significantly higher rates of N and P cycling than Eucalyptus monocultures. It is therefore important to select N 2 -fixing species with readily decomposable litter and high rates of nutrient cycling, as well as high rates of N 2 -fixation. While the dynamics of N 2 -fixation in tree stands are not well understood, it appears as though eucalypts can benefit from fixed N as early as the first or second year following plantation establishment. A meta-analysis of 18 published studies revealed several trials in which mixtures were significantly (P < 0.001) more productive than monocultures, and no instances in which mixtures were less productive than monocultures. Regression analyses of such data were more informative than indices of relative yield, and were more informative in trials that contrasted four or more different species compositions. Thus replacement series examining compositions of 100:0, 67:33, 33:67, and 0:100 were more informative than minimalist 100:0, 50:50 and 0:100 series. #

Forests are critical habitats for biodiversity and they are also essential for the provision of a wide range of ecosystem services that are important to human well-being. There is increasing evidence that biodiversity contributes to forest ecosystem functioning and the provision of ecosystem services. Here we provide a review of forest ecosystem services including biomass production, habitat provisioning services, pollination, seed dispersal, resistance to wind storms, fire regulation and mitigation, pest regulation of native and invading insects, carbon sequestration, and cultural ecosystem services, in relation to forest type, structure and diversity. We also consider relationships between forest biodiversity and multifunctionality, and trade-offs among ecosystem services. We compare the concepts of ecosystem processes, functions and services to clarify their definitions. Our review of published studies indicates a lack of empirical studies that establish quantitative and causal relationships between forest biodiversity and many important ecosystem services. The literature is highly skewed; studies on provisioning of nutrition and energy, and on cultural services, delivered by mixed-species forests are under-represented. Planted forests offer ample opportunity for optimising their composition and diversity because replanting after harvesting is a recurring process. Planting mixed-species forests should be given more consideration as they are likely to provide a wider range of ecosystem services within the forest and for adjacent land uses. This review also serves as the introduction to this special issue of Biodiversity and Conservation on various aspects of forest biodiversity and ecosystem services.

Remote sensing is revolutionizing the way we study forests, and recent technological advances mean we are now able -for the first time -to identify and measure the crown dimensions of individual trees from airborne imagery. Yet to make full use of these data for quantifying forest carbon stocks and dynamics, a new generation of allometric tools which have tree height and crown size at their centre are needed. Here, we compile a global database of 108753 trees for which stem diameter, height and crown diameter have all been measured, including 2395 trees harvested to measure aboveground biomass. Using this database, we develop general allometric models for estimating both the diameter and aboveground biomass of trees from attributes which can be remotely sensed -specifically height and crown diameter. We show that tree height and crown diameter jointly quantify the aboveground biomass of individual trees and find that a single equation predicts stem diameter from these two variables across the world's forests. These new Correspondence: Tommaso Jucker, tel. +44 1223 333911, fax: +44 1223 333953,

Climate models predict an increase in the intensity and frequency of drought episodes in the Northern Hemisphere. Among terrestrial ecosystems, forests will be profoundly impacted by drier climatic conditions, with drastic consequences for the functions and services they supply. Simultaneously, biodiversity is known to support a wide range of forest ecosystem functions and services. However, whether biodiversity also improves the resistance of these ecosystems to drought remains unclear. We compared soil drought exposure levels in a total of 160 forest stands within five major forest types across Europe along a gradient of tree species diversity. We assessed soil drought exposure in each forest stand by calculating the stand-level increase in carbon isotope composition of late wood from a wet to a dry year (Δδ 13 C S ). Δδ 13 C S exhibited a negative linear relationship with tree species diversity in two forest types, suggesting that species interactions in these forests diminished the drought exposure of the ecosystem. However, the other three forest types were unaffected by tree species diversity. We conclude that higher diversity enhances resistance to drought events only in drought-prone environments. Managing forest ecosystems for high tree species diversity does not necessarily assure improved adaptability to the more severe and frequent drought events predicted for the future.climate change | ecology | species interaction | carbon isotope composition | FundivEUROPE

Mixing of complementary tree species may increase stand productivity, mitigate the effects of drought and other risks, and pave the way to forest production systems which may be more resource-use efficient and stable in the face of climate change. However, systematic empirical studies on mixing effects are still missing for many commercially important and widespread species combinations. Here we studied the growth of Scots pine (Pinus sylvestris L.) and European beech (Fagus sylvatica L.) in mixed versus pure stands on 32 triplets located along a productivity gradient through Europe, reaching from Sweden to Bulgaria and from Spain to the Ukraine. Stand inventory and taking increment cores on the mainly 60-80 year-old trees and 0.02-1.55 ha sized, fully stocked plots provided insight how species mixing modifies the structure, dynamics and productivity compared with neighbouring pure stands. In mixture standing volume (?12 %), stand density (?20 %), basal area growth (?12 %), and stand volume growth (?8 %) were higher Communicated by Peter Biber. than the weighted mean of the neighbouring pure stands. Scots pine and European beech contributed rather equally to the overyielding and overdensity. In mixed stands mean diameter (?20 %) and height (?6 %) of Scots pine was ahead, while both diameter and height growth of European beech were behind (-8 %). The overyielding and overdensity were independent of the site index, the stand growth and yield, and climatic variables despite the wide variation in precipitation (520-1175 mm year -1 ), mean annual temperature (6-10.5°C), and the drought index by de Martonne (28-61 mm°C -1 ) on the sites. Therefore, this species combination is potentially useful for increasing productivity across a wide range of site and climatic conditions. Given the significant overyielding of stand basal area growth but the absence of any relationship with site index and climatic variables, we hypothesize that the overyielding and overdensity results from several different types of interactions (light-, water-, and nutrient-related) that are all important in different circumstances. We discuss the relevance of the results for ecological theory and for the ongoing silvicultural transition from pure to mixed stands and their adaptation to climate change. Electronic supplementary material

Biomass and leaf area equations are often required to assess or model forest productivity, carbon stocks and other ecosystem services. These factors are influenced by climate, age and stand structural attributes including stand density and tree species diversity or species composition. However, such covariates are rarely included in biomass and leaf area equations. We reviewed the literature and built a database of biomass and leaf area equations for 24 European tree species and 3 introduced species. The final dataset contained 973 equations. Most of the equations were site-specific and therefore restricted to the edaphic, climatic and stand structural conditions of the given site. To overcome this limitation, the database was used to develop regional species-specific equations that can be used in a wide range of stands and to quantify the effects of climate, age and stand structure on biomass or leaf area. The analysis showed considerable inter-and intra-specific variability in biomass relationships. The intra-specific variability was related to climate, age or stand characteristics, while the inter-specific variability was correlated with traits such as wood density, specific leaf area and shade tolerance. The analysis also showed that foliage mass is more variable than stem or total aboveground biomass, both within and between species, and these biomass components have contrasting responses to age and changes in stand structure. Despite the large number of published equations, many species are still not well represented. Therefore, generic equations were developed that include species-specific wood density instead of species identity. Further improvements may be possible if future studies quantify the stand structure of individual tree neighbourhoods instead of using the stand means for all trees sampled with the given stand.