Phosphorus (P) is a key limiting nutrient in subtropical forests and mixed forests with broadleaved species have been expected to stimulate P cycling, compared to pure conifer plantations. However, the mixture effect of Chinese fir (Cunninghamia lanceolata (Lamb.) Hook.) and broadleaved species on rhizosphere soil and coniferous tree P dynamics is unclear. In our study, eight plots of a single species of a Chinese fir plantation (pure plantation, PP) and eight mixed plantations (mixed plantation, MP) with broadleaved tree species (Michelia macclurei Dandy in Hunan Province or Schima superba Gardn. et Champ. in Fujian Province) were selected in subtropical China. Six P fractions in the rhizosphere and bulk soils were analyzed by a modified Hedley P fractionation method. Phosphorus fractions and nitrogen (N) concentrations in different root orders, different age fresh needles and twigs, and needle and twig litter of Chinese fir were measured. Our results showed that available P, slowly released P, occluded P, and the total extractable P in rhizosphere soil were significantly higher in MP than PP (p < 0.05). In contrast, P and N concentrations in the transportive roots and two-year old needles were generally higher in PP than MP. Meanwhile, the slowly released P, occluded P, total extractable P, and residual P in rhizosphere soil were negatively correlated with P concentrations in young (absorptive and transportive roots, one- and two-year old needles) but not old tissues (storative roots, three-year old needles and litters). In conclusion, mixture may increase soil P availability through the rhizosphere effect, but can decrease P and N concentration of Chinese fir tissues by competition between Chinese fir and broadleaved species. Clearly, the mixture effect may differ in soil and plant nutrients, and this issue needs be taken into consideration when converting a pure conifer plantation into a mixed-species forest.

Species enter and persist in local communities because of their ecological fit to local conditions, and recently, ecologists have moved from measuring diversity as species richness and evenness, to using measures that reflect species ecological differences. There are two principal approaches for quantifying species ecological differences: functional (trait-based) and phylogenetic pairwise distances between species. Both approaches have produced new ecological insights, yet at the same time methodological issues and assumptions limit them. Traits and phylogeny may provide different, and perhaps complementary, information about species' differences. To adequately test assembly hypotheses, a framework integrating the information provided by traits and phylogenies is required. We propose an intuitive measure for combining functional and phylogenetic pairwise distances, which provides a useful way to assess how functional and phylogenetic distances contribute to understanding patterns of community assembly. Here, we show that both traits and phylogeny inform community assembly patterns in alpine plant communities across an elevation gradient, because they represent complementary information. Differences in historical selection pressures have produced variation in the strength of the trait-phylogeny correlation, and as such, integrating traits and phylogeny can enhance the ability to detect assembly patterns across habitats or environmental gradients. © 2013 John Wiley & Sons Ltd/CNRS.

Accelerating rates of species extinction have prompted a growing number of researchers to manipulate the richness of various groups of organisms and examine how this aspect of diversity impacts ecological processes that control the functioning of ecosystems. We summarize the results of 44 experiments that have manipulated the richness of plants to examine how plant diversity affects the production of biomass. We show that mixtures of species produce an average of 1.7 times more biomass than species monocultures and are more productive than the average monoculture in 79% of all experiments. However, in only 12% of all experiments do diverse polycultures achieve greater biomass than their single most productive species. Previously, a positive net effect of diversity that is no greater than the most productive species has been interpreted as evidence for selection effects, which occur when diversity maximizes the chance that highly productive species will be included in and ultimately dominate the biomass of polycultures. Contrary to this, we show that although productive species do indeed contribute to diversity effects, these contributions are equaled or exceeded by species complementarity, where biomass is augmented by biological processes that involve multiple species. Importantly, both the net effect of diversity and the probability of polycultures being more productive than their most productive species increases through time, because the magnitude of complementarity increases as experiments are run longer. Our results suggest that experiments to date have, if anything, underestimated the impacts of species extinction on the productivity of ecosystems.

Numerous grassland experiments have found evidence for a complementarity effect, an increase in productivity with higher plant species richness due to niche partitioning. However, empirical tests of complementarity in natural forests are rare. We conducted a spatially explicit analysis of 518 433 growth records for 274 species from a 50-ha tropical forest plot to test neighborhood complementarity, the idea that a tree grows faster when it is surrounded by more dissimilar neighbors. We found evidence for complementarity: focal tree growth rates increased by 39.8% and 34.2% with a doubling of neighborhood multi-trait dissimilarity and phylogenetic dissimilarity, respectively. Dissimilarity from neighbors in maximum height had the most important effect on tree growth among the six traits examined, and indeed, its effect trended much larger than that of the multitrait dissimilarity index. Neighborhood complementarity effects were strongest for light-demanding species, and decreased in importance with increasing shade tolerance of the focal individuals. Simulations demonstrated that the observed neighborhood complementarities were sufficient to produce positive stand-level biodiversity-productivity relationships. We conclude that neighborhood complementarity is important for productivity in this tropical forest, and that scaling down to individual-level processes can advance our understanding of the mechanisms underlying stand-level biodiversity-productivity relationships.

Forest diversity-productivity relationships have been intensively investigated in recent decades. However, few studies have considered the interplay between species and structural diversity in driving productivity. We analyzed these factors using data from 52 permanent plots in southwestern Germany with more than 53,000 repeated tree measurements. We used basal area increment as a proxy for productivity and hypothesized that: (1) structural diversity would increase tree and stand productivity, (2) diversity-productivity relationships would be weaker for species diversity than for structural diversity, and (3) species diversity would also indirectly impact stand productivity via changes in size structure. We measured diversity using distance-independent indices. We fitted separate linear mixed-effects models for fir, spruce and beech at the tree level, whereas at the stand level we pooled all available data. We tested our third hypothesis using structural equation modeling. Structural and species diversity acted as direct and independent drivers of stand productivity, with structural diversity being a slightly better predictor. Structural diversity, but not species diversity, had a significant, albeit asymmetric, effect on tree productivity. The functioning of structurally diverse, mixed forests is influenced by both structural and species diversity. These sources of trait diversity contribute to increased vertical stratification and crown plasticity, which in turn diminish competitive interferences and lead to more densely packed canopies per unit area. Our research highlights the positive effects of species diversity and structural diversity on forest productivity and ecosystem dynamics.

Theory suggests that plant interactions at the neighbourhood scale play a fundamental role in regulating biodiversity-productivity relationships (BPRs) in tree communities. However, empirical evidence of this prediction is rare, as little is known about how neighbourhood interactions scale up to influence community BPRs. Here, using a biodiversity-ecosystem functioning experiment, we provide insights into processes underlying BPRs by demonstrating that diversity-mediated interactions among local neighbours are a strong regulator of productivity in species mixtures. Our results show that local neighbourhood interactions explain over half of the variation in observed community productivity along a diversity gradient. Overall, individual tree growth increased with neighbourhood species richness, leading to a positive BPR at the community scale. The importance of local-scale neighbourhood effects for regulating community productivity, however, distinctly increased with increasing community species richness. Preserving tree species diversity at the local neighbourhood scale, thus seems to be a promising way for promoting forest productivity.

Studies on tree communities have demonstrated that species diversity can enhance forest productivity, but the driving mechanisms at the local neighbourhood level remain poorly understood. Here, we use data from a large-scale biodiversity experiment with 24 subtropical tree species to show that neighbourhood tree species richness generally promotes individual tree productivity. We found that the underlying mechanisms depend on a focal tree's functional traits: For species with a conservative resource-use strategy diversity effects were brought about by facilitation, and for species with acquisitive traits by competitive reduction. Moreover, positive diversity effects were strongest under low competition intensity (quantified as the total basal area of neighbours) for acquisitive species, and under high competition intensity for conservative species. Our findings demonstrate that net biodiversity effects in tree communities can vary over small spatial scales, emphasising the need to consider variation in local neighbourhood interactions to better understand effects at the community level.© 2017 John Wiley & Sons Ltd/CNRS.

We investigated the relationship between plant diversity and ecological function (production and nutrient cycling) in tropical tree plantations. Old plantations (65-72 years) of four different species, namely Araucaria cunninghamii, Agathis robusta, Toona ciliata and Flindersia brayleyana, as well as natural secondary forest were examined at Wongabel State Forest, in the wet tropics region of Queensland, Australia. Two young plantations (23 years) of Araucaria cunninghamii and Pinus caribaea were also examined. The close proximity of the older plantations and natural forests meant they had similar edaphic and climatic conditions. All plantations had been established as monocultures, but had been colonised by a range of native woody plants from the nearby rainforest. The extent to which this had occurred varied with the identity of the plantation species (from 2 to 17 species in 0.1 ha blocks). In many cases these additional species had grown up and joined the forest canopy. This study is one of the few to find a negative relationship between overstorey plant diversity and productivity. The conversion of natural forest with highly productive, low-diversity gymnosperm-dominated plantations (young and old Araucaria cunninghamii and Pinus caribaea) was found to be associated with lower soil nutrient availability (approximately five times less phosphorus and 2.5 times less nitrogen) and lower soil pH (mean=6.28) compared to the other, less productive plantations. The dominant effects of two species, Araucaria cunninghamii and Hodgkinsonia frutescens, indicate that ecosystem functions such as production and nutrient availability are not determined solely by the number of species, but are more likely to be determined by the characteristics of the species present. This suggests that monoculture plantations can be used to successfully restore some functions (e.g. nutrient cycling and production), but that the level to which such functions can be restored will depend upon the species chosen and site conditions.

As distinct community assembly processes can produce similar community patterns, assessing the ecological mechanisms promoting coexistence in hyperdiverse rainforests remains a considerable challenge. We use spatially explicit neighbourhood models of tree growth to quantify how functional trait and phylogenetic similarities predict variation in growth and crowding effects for the 315 most abundant tree species in a 25-ha lowland rainforest plot in Ecuador. We find that functional trait differences reflect variation in (1) species maximum potential growth, (2) the intensity of interspecific interactions for some species, and (3) species sensitivity to neighbours. We find that neighbours influenced tree growth in 28% of the 315 focal tree species. Neighbourhood effects are not detected in the remaining 72%, which may reflect the low statistical power to model rare taxa and/or species insensitivity to neighbours. Our results highlight the spectrum of ways in which functional trait differences can shape community dynamics in highly diverse rainforests.© 2016 John Wiley & Sons Ltd/CNRS.

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.Copyright © 2018 The Authors, some rights reserved; exclusive licensee American Association for the Advancement of Science. No claim to original U.S. Government Works.

细根在森林生态系统C分配和养分循环过程中发挥着重要作用, 但对地下细根与植物多样性之间关系的研究相对较少。该研究选择中亚热带从单一树种的杉木(Cunninghamia lanceolata)人工林到多树种的常绿阔叶林(青冈(Cyclobalanopsis glauca)-石栎(Lithocarpus glaber)林)的不同植物多样性梯度, 用根钻法采集细根并测定其生物量, 用Win-RHIZO 2005C根系分析系统测定细根形态参数, 以验证以下3个假设: 1)植物种类丰富度高的林分其细根生产存在&ldquo;地下超产&rdquo;现象; 2)根系空间生态位的分离水平是否随着植物多样性增多而增大? 3)细根是否通过形态可塑性对林木竞争做出响应？结果显示: 从单一树种的杉木人工林到植物种类较复杂的青冈-石栎常绿阔叶林, 0&ndash;30 cm土层的林分细根总生物量和活细根生物量均呈增加的趋势, 即细根总生物量为杉木林(305.20 g&middot;m^&ndash;2) Pinus massoniana)林(374.25 g&middot;m^&ndash;2) Choerospondias axillaris)林(537.42 g&middot;m^&ndash;2) &ndash;2</sup>), 活细根生物量为杉木林(268.74 g&middot;m^&ndash;2) &ndash;2</sup>) &ndash;2</sup>) &ndash;2</sup>), 各森林类型之间的细根总生物量差异显著(p

The relationship between plant species diversity and primary productivity has long been a hot topic in biodiversity research. We examined the relationship between species richness and productivity in four typical grasslands of northern China at different spatial scales. At the community scale, a positive correlation was found for six of seven communities, and a unimodal pattern was found only for one community (<i>Stipa glareosa community</i>). At large scale (vegetation type or landscape/region), a significant positive relationship was found. Species richness ranged from 4 to 35 species, and community above-ground productivity from 13 to 368 g·m^–2·yr^–1. The highest species richness and above-ground productivity were found in the alpine meadow community, followed by meadow steppe, typical steppe and desert steppe.

利用2002–2004年内蒙古和甘肃南部几种典型草地的实测资料, 研究了不同尺度物种丰富度与生产力的关系, 并初步探讨了其形成机制。结果显示, 温带草地的物种丰富度随生产力的增加而增加, 但受空间尺度影响。在群落尺度(同一群落), 在7种样方数大于15的群落中, 仅沙生针茅(Stipa glareosa)群落物种丰富度与生产力呈现单峰型关系, 其余均呈现线性正相关关系; 在植被类型尺度, 物种丰富度–生产力之间表现为显著的正相关关系; 在研究区尺度, 物种丰富度随生产力的增加而显著增加。研究还表明, 研究区群落生产力的变化范围为13–368 g·m^–2·yr^–1, 物种丰富度为4–35 种; 生产力从高到低的顺序为: 高寒草甸>草甸草原>典型草原>荒漠草原。

Though many processes are involved in determining which species coexist and assemble into communities, competition is among the best studied. One hypothesis about competition's contribution to community assembly is that more closely related species are less likely to coexist. Though empirical evidence for this hypothesis is mixed, it remains a common assumption in certain phylogenetic approaches for inferring the effects of environmental filtering and competitive exclusion. Here, we relate modern coexistence theory to phylogenetic community assembly approaches to refine expectations for how species relatedness influences the outcome of competition. We argue that two types of species differences determine competitive exclusion with opposing effects on relatedness patterns. Importantly, this means that competition can sometimes eliminate more different and less related taxa, even when the traits underlying the relevant species differences are phylogenetically conserved. Our argument leads to a reinterpretation of the assembly processes inferred from community phylogenetic structure.

The importance of interspecific competition as a cause of resource partitioning among species has been widely assumed but rarely tested. Using neighbor removals in combination with 15N tracer additions in the field, we examined variation among three alpine species in the uptake of 15N-NH4+, 15N-NO3-, and 15N-13C-[2]-glycine in intact neighborhoods, when paired with a specific neighbor, and when all neighbors were removed. Species varied in the capacity to take up 15N-labeled NH4+, NO3-, and glycine in intact neighborhoods and in interspecific pairs. When interspecific neighbor pairs were compared with no neighbor controls, neighbors reduced 15N uptake in target species by as much as 50%, indicating competition for N. Furthermore, neighbor identity influenced the capacity of species to take up different forms of N. Thus, competition within interspecific neighbor pairs often caused reduced uptake of a particular form of N, as well as shifts to uptake of an alternative form of N. Such shifts in resource use as a result of competition are an implicit assumption in studies of resource partitioning but have rarely been documented. Our study suggests that plasticity in the uptake of different forms of N may be a mechanism by which cooccurring plants reduce competition for N.

Negative density dependence (NDD) and environmental filtering (EF) shape community assembly, but their relative importance is poorly understood. Recent studies have shown that seedling's mortality risk is positively related to the phylogenetic relatedness of neighbours. However, natural enemies, whose depredations often cause NDD, respond to functional traits of hosts rather than phylogenetic relatedness per se. To understand the roles of NDD and EF in community assembly, we assessed the effects on seedling mortality of functional similarity, phylogenetic relatedness and stem density of neighbouring seedlings and adults in a species-rich tropical forest. Mortality risks increased for common species when their functional traits departed substantially from the neighbourhood mean, and for all species when surrounded by close relatives. This indicates that NDD affects community assembly more broadly than does EF, and leads to the tentative conclusion that natural enemies respond to phylogenetically correlated traits. Our results affirm the prominence of NDD in structuring species-rich communities.© 2011 Blackwell Publishing Ltd/CNRS.

Recent studies on species coexistence suggest that density dependence is an important mechanism regulating plant populations. However, there have been few studies of density dependence conducted for more than one life-history stage or that control for habitat heterogeneity, which may influence spatial patterns of survival and mask density dependence. We explored the prevalence of density dependence across multiple life stages, and the effects of controlling for habitat heterogeneity, in a temperate forest in northeast China. We used generalized linear mixed-effects models to test for density-dependent mortality of seedlings and spatial point pattern analysis to detect density dependence for sapling-to-juvenile transitions. Conspecific neighbors had a negative effect on survival of plants in both life stages. At the seedling stage, we found a negative effect of conspecific seedling neighbors on survival when analyzing all species combined. However, in species-level analyses, only 2 of 11 focal species were negatively impacted by conspecific neighbors, indicating wide variation among species in the strength of density dependence. Controlling for habitat heterogeneity did not alter our findings of density dependence at the seedling stage. For the sapling-to-juvenile transition stage, 11 of 15 focal species showed patterns of local scale (<10 m) conspecific thinning, consistent with negative density dependence. The results varied depending on whether we controlled for habitat heterogeneity, indicating that a failure to account for habitat heterogeneity can obscure patterns of density dependence. We conclude that density dependence may promote tree species coexistence by acting across multiple life-history stages in this temperate forest.

An assemblage of tree species with different crown properties creates heterogeneous environments at the canopy level. Changes of functional leaf traits are expected, especially those related to light interception and photosynthesis. Chlorophyll a fluorescence (ChlF) properties in dark-adapted leaves, specific leaf area, leaf nitrogen content (N) and carbon isotope composition (δ13C) were measured on Picea abies (L.) H.Karst., Pinus sylvestris L. and Betula pendula Roth. in monospecific and mixed boreal forests in Europe, in order to test whether they were affected by stand species richness and composition. Photosynthetic efficiency, assessed by induced emission of leaf ChlF, was positively influenced in B. pendula by species richness, whereas P. abies showed higher photosynthetic efficiency in monospecific stands. Pinus sylvestris had different responses when it coexisted with P. abies or B. pendula. The presence of B. pendula, but not of P. abies, in the forest had a positive effect on the efficiency of photosynthetic electron transport and N in P. sylvestris needles, and the photosynthetic responses were positively correlated with an increase of leaf δ13C. These effects on P. sylvestris may be related to high light availability at the canopy level due to the less dense canopy of B. pendula. The different light requirements of coexisting species was the most important factor affecting the distribution of foliage in the canopy, driving the physiological responses of the mixed species. Future research directions claim to enhance the informative potential of the methods to analyse the responses of pure and mixed forests to environmental factors, including a broader set of plant species' functional traits and physiological responses.© The Author 2017. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.

Plant diversity generally promotes biomass production, but how the shape of the response curve changes with time remains unclear. This is a critical knowledge gap because the shape of this relationship indicates the extent to which loss of the first few species will influence biomass production. Using two long-term (≥13 years) biodiversity experiments, we show that the effects of diversity on biomass productivity increased and became less saturating over time. Our analyses suggest that effects of diversity-dependent ecosystem feedbacks and interspecific complementarity accumulate over time, causing high-diversity species combinations that appeared functionally redundant during early years to become more functionally unique through time. Consequently, simplification of diverse ecosystems will likely have greater negative impacts on ecosystem functioning than has been suggested by short-term experiments.

Productivity of tree plantations is a function of the supply, capture and efficiency of use of resources, as outlined in the Production Ecology Equation. Species interactions in mixed-species stands can influence each of these variables. The importance of resource-use efficiency in determining forest productivity has been clearly demonstrated in monocultures; however, substantial knowledge gaps remain for mixtures. This review examines how the physiology and morphology of a given species can vary depending on whether it grows in a mixture or monoculture. We outline how physiological and morphological shifts within species, resulting from interactions in mixtures, may influence the three variables of the Production Ecology Equation, with an emphasis on nutrient resources [nitrogen (N) and phosphorus (P)]. These include (i) resource availability, including soil nutrient mineralization, N₂ fixation and litter decomposition; (ii) proportion of resources captured, resulting from shifts in spatial, temporal and chemical patterns of root dynamics; (iii) resource-use efficiency. We found that more than 50% of mixed-species studies report a shift to greater above-ground nutrient content of species grown in mixtures compared to monocultures, indicating an increase in the proportion of resources captured from a site. Secondly, a meta-analysis showed that foliar N concentrations significantly increased for a given species in a mixture containing N₂-fixing species, compared to a monoculture, suggesting higher rates of photosynthesis and greater resource-use efficiency. Significant shifts in N- and P-use efficiencies of a given species, when grown in a mixture compared to a monoculture, occurred in over 65% of studies where resource-use efficiency could be calculated. Such shifts can result from changes in canopy photosynthetic capacities, changes in carbon allocation or changes to foliar nutrient residence times of species in a mixture. We recommend that future research focus on individual species' changes, particularly with respect to resource-use efficiency (including nutrients, water and light), when trees are grown in mixtures compared to monocultures. A better understanding of processes responsible for changes to tree productivity in mixed-species tree plantations can improve species, and within-species, selection so that the long-term outcome of mixtures is more predictable.

The phosphorus nutrition status of European forests has decreased significantly in recent decades. For a deeper understanding of complementarity and competition in terms of P acquisition in temperate forests, we have analyzed α-diversity, organic layer and mineral soil P, P nutrition status, and different concepts of P use efficiency (PUE) in Fagus sylvatica L. (European beech) and Picea abies (L.) H. Karst. (Norway spruce). Using a subset of the Second National Soil Survey in Germany, we correlated available data on P in the organic layer and soil with α-diversity indices for beech and spruce forests overall and for individual vegetation layers (tree, shrub, herb, and moss layers). Moreover, we investigated α-diversity feedbacks on P nutrition status and PUE of both tree species. The overall diversity of both forest ecosystems was largely positively related to P content in the organic layer and soil, but there were differences among the vegetation layers. Diversity in the tree layer of both forest ecosystems was negatively related to the organic layer and soil P. By contrast, shrub diversity showed no correlation to P, while herb layer diversity was negatively related to P in the organic layer but positively to P in soil. A higher tree layer diversity was slightly related to increased P recycling efficiency (PPlant/Porganic layer) in European beech and P uptake efficiency (PPlant/Psoil) in Norway spruce. The diversity in the herb layer was negatively related to P recycling and uptake efficiency in European beech and slightly related to P uptake efficiency in Norway spruce. In spruce forests, overall and herb species richness led to significantly improved tree nutrition status. Our results confirm significant, non-universal relationships between P and diversity in temperate forests with variations among forest ecosystems, vegetation layers, and P in the organic layer or soil. In particular, tree species diversity may enhance complementarity and hence also P nutrition of dominant forest trees through higher PUE, whereas moss and herb layers seemed to show competitive relationships among each other in nutrient cycling.

Plant species use different strategies for maximizing growth and fitness under changing environmental conditions. At the ecosystem level, seedlings in particular compete with other vegetation components for light and nitrogen (N), which often constitute growth-limiting resources. In this study, we investigated the effect of light availability on the competition for N between seedlings of European beech and sycamore maple and analysed the consequences of this competition for the composition of N metabolites in fine roots. Our results show different strategies in N acquisition between beech and sycamore maple. Both species responded to reduced light availability by adapting their morphological and physiological traits with a decrease in biomass and net assimilation rate and an increase in specific leaf area and leaf area ratio. For beech seedlings, competition with sycamore maple led to a reduction in organic N uptake capacity. Reduced light availability led to a decrease in ammonium, but an increase in glutamine-N uptake capacity in sycamore maple. However, this response was stronger compared with that of beech and was accompanied by reduced growth. Thus, our results suggest better adaptation of N acquisition to reduced light availability in beech compared with sycamore maple seedlings.

To investigate the short-term consequences of direct competition between beech and sycamore maple on root N uptake and N composition, mycorrhizal seedlings of both tree species were incubated for 4 days (i.e. beech only, sycamore maple only or both together) in an artificial nutrient solution with low N availability. On the fourth day, N uptake experiments were conducted to study the effects of competition on inorganic and organic N uptake. For this purpose, multiple N sources were applied with a single label. Furthermore, fine roots were sampled and analysed for total amino acids, soluble protein, total nitrogen, nitrate and ammonium content. Our results clearly show that both tree species were able to use inorganic and organic N sources. Uptake of inorganic and organic N by beech roots was negatively affected in the presence of the competing tree species. In contrast, the presence of beech stimulated inorganic N uptake by sycamore maple roots. Both the negative effect of sycamore maple on N uptake of beech and the positive effect of beech on N uptake of sycamore maple led to an increase in root soluble protein in beech, despite an overall decrease in total N concentration. Thus, beech compensated for the negative effects of the tree competitor on N uptake by incorporating less N into structural N components, but otherwise exhibited the same strategy as the competitor, namely, enhancing soluble protein levels in roots when grown under competition. It is speculated that enhanced enzyme activities of so far unknown nature are required in beech as a defence response to inter-specific competition.

Agricultural landscape homogenization has detrimental effects on biodiversity and key ecosystem services. Increasing agricultural landscape heterogeneity by increasing seminatural cover can help to mitigate biodiversity loss. However, the amount of seminatural cover is generally low and difficult to increase in many intensively managed agricultural landscapes. We hypothesized that increasing the heterogeneity of the crop mosaic itself (hereafter "crop heterogeneity") can also have positive effects on biodiversity. In 8 contrasting regions of Europe and North America, we selected 435 landscapes along independent gradients of crop diversity and mean field size. Within each landscape, we selected 3 sampling sites in 1, 2, or 3 crop types. We sampled 7 taxa (plants, bees, butterflies, hoverflies, carabids, spiders, and birds) and calculated a synthetic index of multitrophic diversity at the landscape level. Increasing crop heterogeneity was more beneficial for multitrophic diversity than increasing seminatural cover. For instance, the effect of decreasing mean field size from 5 to 2.8 ha was as strong as the effect of increasing seminatural cover from 0.5 to 11%. Decreasing mean field size benefited multitrophic diversity even in the absence of seminatural vegetation between fields. Increasing the number of crop types sampled had a positive effect on landscape-level multitrophic diversity. However, the effect of increasing crop diversity in the landscape surrounding fields sampled depended on the amount of seminatural cover. Our study provides large-scale, multitrophic, cross-regional evidence that increasing crop heterogeneity can be an effective way to increase biodiversity in agricultural landscapes without taking land out of agricultural production.

Plant diversity and niche complementarity had progressively stronger effects on ecosystem functioning during a 7-year experiment, with 16-species plots attaining 2.7 times greater biomass than monocultures. Diversity effects were neither transients nor explained solely by a few productive or unviable species. Rather, many higher-diversity plots outperformed the best monoculture. These results help resolve debate over biodiversity and ecosystem functioning, show effects at higher than expected diversity levels, and demonstrate, for these ecosystems, that even the best-chosen monocultures cannot achieve greater productivity or carbon stores than higher-diversity sites.

Two main effects are proposed to explain biodiversity-ecosystem functioning relationships: niche complementarity and selection effects. Both can be functionally defined using the functional diversity (FD) and functional identity (FI) of the community respectively. Herein, we present results from the first tree diversity experiment that separated the effect of selection from that of complementarity by varying community composition in high-density plots along a gradient of FD, independent of species richness and testing for the effects of FD and community weighted means of traits (a proxy for FI) on stem biomass increment (a proxy for productivity). After 4 years of growth, most mixtures did not differ in productivity from the averages of their respective monocultures, but some did overyield significantly. Those positive diversity effects resulted mostly from selection effects, primarily driven by fast-growing deciduous species and associated traits. Net diversity effect did not increase with time over 4 years. © 2016 John Wiley & Sons Ltd/CNRS.

. Tropical forests play a significant role in the global carbon cycle and global climate. However, tropical carbon cycling and the feedbacks from tropical ecosystems to the climate system remain critical uncertainties in the current generation of carbon–climate models. One of the major uncertainties comes from the lack of representation of phosphorus (P), currently believed to be the most limiting nutrient in tropical regions. Here we introduce P dynamics and C–N–P interactions into the CLM4-CN (Community Land Model version 4 with prognostic Carbon and Nitrogen) model and investigate the role of P cycling in controlling the productivity of tropical ecosystems. The newly developed CLM-CNP model includes all major biological and geochemical processes controlling P availability in soils and the interactions between C, N, and P cycles. Model simulations at sites along a Hawaiian soil chronosequence indicate that the introduction of P limitation greatly improved the model performance at the P-limited site. The model is also able to capture the shift in nutrient limitation along this chronosequence (from N limited to P limited), as shown in the comparison of model-simulated plant responses to fertilization with the observed data. Model simulations at Amazonian forest sites show that CLM-CNP is capable of capturing the overall trend in NPP (net primary production) along the P availability gradient. This comparison also suggests a significant interaction between nutrient limitation and land use history. Model experiments under elevated atmospheric CO2 ([CO2]) conditions suggest that tropical forest responses to increasing [CO2] will interact strongly with changes in the P cycle. We highlight the importance of two feedback pathways (biochemical mineralization and desorption of secondary mineral P) that can significantly affect P availability and determine the extent of P limitation in tropical forests under elevated [CO2]. Field experiments with elevated CO2 are therefore needed to help quantify these important feedbacks. CO2 doubling model experiments show that tropical forest response to elevated [CO2] can only be predicted if the interactions between C cycle and nutrient dynamics are well understood and represented in models. Predictive modeling of C–nutrient interactions will have important implications for the prediction of future carbon uptake and storage in tropical ecosystems and global climate change.\n

通过对福建三明27年生杉木(Cunninghamia lanceolata)-观光木(Tsoongiodendron odorum)混交林（混交比例2：1）及杉木纯林群落N、P养分循环进行为期2年的研究。结果表明，混交林中杉木和观光木地上各组分的N、P含量大小均为叶>活枝（或皮）>枯枝>干，而根系的则随径级的减小而增大，且观光木各组分的N含量均高于杉木的；混交林群落的N、P总积累量达585.223 kg&middot;hm－2和128.784 kg&middot;hm－2，分别是纯林群落的1.5倍和1.3倍。混交林群落N、P养分年归

Deficiency in availability of phosphorus (P) in forest soils of southern China, where Chinese fir (Cunninghamia lanceolata (Lamb.) Hook.) stands have been established, greatly affects Chinese fir productivity. Deploying clones with high P-use efficiency could be a more attractive option than P fertilization to enhance the commercial productivity of Chinese fir. Thus, an understanding of the adaptive responses of roots to spatiotemporal variability of P, which is often ignored in previous studies, is essential. In this study, we report the root responses of a Chinese fir clone with high P-use efficiency to the spatially heterogeneous P availability in soil during various periods of time when P availability was controlled. The results showed significantly (p &lt; 0.05) more root proliferation in the form of root length, root surface area, root volume, and root dry matter accumulation in P-replete than in P-poor patches when the patch persisted for a short period of time (50 days). As the patches persisted longer (100–200 days), root proliferation in P-replete patches appeared to decline. P-replete patches that persisted for short (50 days) and long (200 days) periods resulted in high root P accumulation, whereas P-replete patches that persisted for 100–150 days resulted in lower root P accumulation than other patches. It seems that the observed responses in the roots of the Chinese fir clone to heterogeneous P availability might include a comprehensive adjustment of root growth and intraroot P translocation in patches so as to maintain internal P homeostasis, as well as a trade-off between energy cost and production of new roots when the patch persists longer.