Functional traits composition predict macrophytes community productivity along a water depth gradient in a freshwater lake

Functional trait composition of plant communities has been proposed as a helpful key for understanding the mechanisms of biodiversity effects on ecosystem functioning. In this study, we applied a step‐wise modeling procedure to test the relative effects of taxonomic diversity, functional identity, and functional diversity on macrophytes community productivity along water depth gradient. We sampled 42 plots and 1513 individual plants and measured 16 functional traits and abundance of 17 macrophyte species. Results showed that there was a significant decrease in taxonomic diversity, functional identity (i.e., stem dry mass content, leaf [C] and leaf [N]), and functional diversity (i.e., floating leaf, mean Julian flowering date and rooting depth) with increasing water depth. For the multiple‐trait functional diversity (FD) indices, functional richness decreased, while functional divergence increased with water depth gradient. Macrophyte community productivity was strongly determined by functional trait composition within community, but not significantly affected by taxonomic diversity. Community‐weighted means (CWM) showed a two times higher explanatory power relative to FD indices in determining variations in community productivity. For nine of sixteen traits, CWM and FD showed significant correlations with community productivity, although the strength and direction of those relations depended on selected trait. Furthermore, functional composition in a community affected productivity through either additive or opposite effects of CWM and FD, depending on the particular traits being considered. Our results suggested both mechanisms of mass ratio and niche complementarity can operate simultaneously on variations in community productivity, and considering both CWM and FD would lead to a more profound understanding of traits–productivity relationships.     

Using root traits to understand temporal changes in biodiversity effects in grassland mixtures

Biodiversity–ecosystem functioning (BEF) studies typically show that species richness enhances community biomass, but the underlying mechanisms remain debated. Here, we combine metrics from BEF research that distinguish the contribution of dominant species (selection effects, SE) from those due to positive interactions such as resource partitioning (complementarity effects, CE) with a functional trait approach in an attempt to reveal the functional characteristics of species that drive community biomass in species mixtures. In a biodiversity experiment with 16 plant species in monocultures, 4‐species and 16‐species mixtures, we used aboveground biomass to determine the relative contributions of CE and SE to biomass production in mixtures in the second, dry year of the experiment. We also measured root traits (specific root length, root length density, root tissue density and the deep root fraction) of each species in monocultures and linked the calculated community weighted mean (CWM) trait values and trait diversity of mixtures to CE and SE. In the second year of the experiment, community biomass, CE and SE increased compared to the first year. The contribution of SE to this positive effect was greater than that of CE. The increased contribution of SE was associated with root traits: SE increased most in communities with high abundance of species with deep, thick and dense roots. In contrast, changes in CE were not related to trait diversity or CWM trait values. Together, these results suggest that increased positive effects of species richness on community biomass in a dry year were mainly driven by increased dominance of deep‐rooting species, supporting the insurance hypothesis of biodiversity. Positive CE indicates that other positive interactions did occur, but we could not find evidence that belowground resource partitioning or facilitation via root trait diversity was important for community productivity in our biodiversity experiment.     

Leaf decomposition depends on nutritional trait values but increasing trait variability does not always increase decomposition efficiency

The mass ratio (MRH) and niche complementarity (NCH) hypotheses can explain how leaf trait composition drives decomposition, an ecosystem process linked to nutrient cycling and carbon sequestration. However, few studies have used an experiment designed to disentangle the role of the mechanisms proposed by these hypotheses. This is especially true regarding the role of leaf functional traits for decomposition rates in tropical ecosystems. Here, we quantified the biomass loss of 120 leaf mixtures assembled according to four quasi-orthogonal combinations of different mean trait values (community-weighted mean; CWM) and trait variability (functional diversity; FD) of three leaf functional traits (leaf nitrogen and leaf magnesium concentrations and specific leaf area). We found that CWM values of leaf nutritional traits were positively related to greater biomass loss. This supports the hypothesis that the mean trait values of leaf mixtures can drive biomass loss (MRH). However, contrary to NCH expectations, in some circumstances, increasing trait variability of leaf nutritional traits decreased biomass loss. Our results reinforce some previous evidence that, together, CWM and FD can explain leaf decomposition and highlight that the mean resource quality of leaf mixtures is a driver of biomass loss. Also, as previously reported for temperate ecosystems, trait variability does not always increase leaf decomposition in tropical ecosystems. Therefore, there is a need to consider simultaneously both MRH and NCH in future studies, using an appropriate design, keeping in mind that both mechanisms will always be present in any species mixture or combination.     

C,N and P fertilization in an Amazonian rainforest supports stoichiometric dissimilarity as a driver of litter diversity effects on decomposition

Plant leaf litter generally decomposes faster as a group of different species than when individual species decompose alone, but underlying mechanisms of these diversity effects remain poorly understood. Because resource C : N : P stoichiometry (i.e. the ratios of these key elements) exhibits strong control on consumers, we supposed that stoichiometric dissimilarity of litter mixtures (i.e. the divergence in C : N : P ratios among species) improves resource complementarity to decomposers leading to faster mixture decomposition. We tested this hypothesis with: (i) a wide range of leaf litter mixtures of neotropical tree species varying in C : N : P dissimilarity, and (ii) a nutrient addition experiment (C, N and P) to create stoichiometric similarity. Litter mixtures decomposed in the field using two different types of litterbags allowing or preventing access to soil fauna. Litter mixture mass loss was higher than expected from species decomposing singly, especially in presence of soil fauna. With fauna, synergistic litter mixture effects increased with increasing stoichiometric dissimilarity of litter mixtures and this positive relationship disappeared with fertilizer addition. Our results indicate that litter stoichiometric dissimilarity drives mixture effects via the nutritional requirements of soil fauna. Incorporating ecological stoichiometry in biodiversity research allows refinement of the underlying mechanisms of how changing biodiversity affects ecosystem functioning.     

Functional diversity in root and above-ground traits in a fertile grassland shows a detrimental effect on productivity

Plant community functional structure may drive ecosystem functions in relation with (i) the trait values characterizing dominant species, according to the “biomass ratio hypothesis” proposed by Grime, and (ii) thanks to trait dissimilarity among species, according to the “diversity hypothesis” proposed by Tilman. Both propositions have already yielded support but their relative importance and how they impact biomass production in field situations is still not well known. This study therefore tested (i) whether or not there was a close relationship between biomass production and the community-weighted mean trait values (CWM), as expected from the “biomass ratio hypothesis”, and (ii) the impact of the functional diversity (FD_Q) on biomass production, which is expected to be positive according to the “diversity-hypothesis”. The study considered a range of plant assemblages occurring in a wet grassland and five above-ground and four below-ground plant traits were measured to characterize their functional structure. The effects of species diversity, soil water content, soil nitrogen availability and grazing intensity on biomass production were also determined.We showed that biomass production was not related either to species richness and diversity or to any of the resource and disturbance parameters considered. Conversely, the functional structure was found to explain up to 55% of the variability of the biomass production. The results obtained clearly supported the “biomass-ratio hypothesis”. Functional diversity was mainly found to negatively impact biomass production with only poor support to the “diversity hypothesis”. We suggest that such a dilution effect of increasing FD_Q on community primary production may be typical of fertile habitats.In order to significantly improve our understanding of the relationship between functional diversity and ecosystem processes, further studies should consider plant assemblages that have been shaped over the long term and habitats across a wide range of productivity.     

The essential role of biodiversity in the key axes of ecosystem function

Biodiversity is essential for maintaining the terrestrial ecosystem multifunctionality (EMF). Recent studies have revealed that the variations in terrestrial ecosystem functions are captured by three key axes: the maximum productivity, water use efficiency, and carbon use efficiency of the ecosystem. However, the role of biodiversity in supporting these three key axes has not yet been explored. In this study, we combined the (i) data collected from more than 840 vegetation plots across a large climatic gradient in China using standard protocols, (ii) data on plant traits and phylogenetic information for more than 2,500 plant species, and (iii) soil nutrient data measured in each plot. These data were used to systematically assess the contribution of environmental factors, species richness, functional and phylogenetic diversity, and community-weighted mean (CWM) and ecosystem traits (i.e., traits intensity normalized per unit land area) to EMF via hierarchical partitioning and Bayesian structural equation modeling. Multiple biodiversity attributes accounted for 70% of the influence of all the variables on EMF, and ecosystems with high functional diversity had high resource use efficiency. Our study is the first to systematically explore the role of different biodiversity attributes, including species richness, phylogenetic and functional diversity, and CWM and ecosystem traits, in the key axes of ecosystem functions. Our findings underscore that biodiversity conservation is critical for sustaining EMF and ultimately ensuring human well-being.     

Functional diversity (FD), species richness and community composition

Functional diversity is an important component of biodiversity, yet in comparison to taxonomic diversity, methods of quantifying functional diversity are less well developed. Here, we propose a means for quantifying functional diversity that may be particularly useful for determining how functional diversity is related to ecosystem functioning. This measure of functional diversity “FD” is defined as the total branch length of a functional dendrogram. Various characteristics of FD make it preferable to other measures of functional diversity, such as the number of functional groups in a community. Simulating species' trait values illustrates how the relative importance of richness and composition for FD depends on the effective dimensionality of the trait space in which species separate. Fewer dimensions increase the importance of community composition and functional redundancy. More dimensions increase the importance of species richness and decreases functional redundancy. Clumping of species in trait space increases the relative importance of community composition. Five natural communities show remarkably similar relationships between FD and species richness.     

The quest for trait convergence and divergence in community assembly: are null‐models the magic wand?

The relevance of neutral versus niche‐based community assembly rules (i.e. the processes sorting species present in a larger geographical region into local communities) remains to be demonstrated in ecology and biogeography. To attempt to do this, a number of complex null models are increasingly being used that compare observed community functional diversity (FD, i.e. the extent of trait dissimilarity between coexisting species) with randomly simulated FD. However, little is known about the performance of these null models in detecting non‐neutral community assembly rules such as trait convergence and divergence of communities (supposedly revealing habitat selection and limiting similarity, respectively). Here, using both simulated and field communities, I show that assembly rule detection varies systematically with the magnitude of the observed FD, so that these null models do not really succeed in breaking down the observed functional relationships between species. This is a particular concern, making detection of community assembly dependent on: (1) the pool of samples considered, and (2) the capacity of observed FD to correctly discriminate these rules. Null models should be more thoroughly described and validated before being considered as a magic wand to reveal assembly patterns.     

Do temperate tree species diversity and identity influence soil microbial community function and composition?

Studies of biodiversity–ecosystem function in treed ecosystems have generally focused on aboveground functions. This study investigates intertrophic links between tree diversity and soil microbial community function and composition. We examined how microbial communities in surface mineral soil responded to experimental gradients of tree species richness (SR ), functional diversity (FD ), community‐weighted mean trait value (CWM ), and tree identity. The site was a 4‐year‐old common garden experiment near Montreal, Canada, consisting of deciduous and evergreen tree species mixtures. Microbial community composition, community‐level physiological profiles, and respiration were evaluated using phospholipid fatty acid (PLFA ) analysis and the MicroResp^? system, respectively. The relationship between tree species richness and glucose‐induced respiration (GIR ), basal respiration (BR ), metabolic quotient (qCO ₂) followed a positive but saturating shape. Microbial communities associated with species mixtures were more active (basal respiration [BR ]), with higher biomass (glucose‐induced respiration [GIR ]), and used a greater number of carbon sources than monocultures. Communities associated with deciduous tree species used a greater number of carbon sources than those associated with evergreen species, suggesting a greater soil carbon storage capacity. There were no differences in microbial composition (PLFA ) between monocultures and SR mixtures. The FD and the CWM of several functional traits affected both BR and GIR . In general, the CWM of traits had stronger effects than did FD , suggesting that certain traits of dominant species have more effect on ecosystem processes than does FD . Both the functions of GIR and BR were positively related to aboveground tree community productivity. Both tree diversity (SR ) and identity (species and functional identity—leaf habit) affected soil microbial community respiration, biomass, and composition. For the first time, we identified functional traits related to life‐history strategy, as well as root traits that influence another trophic level, soil microbial community function, via effects on BR and GIR .     

Functional dissimilarity across trophic levels as a driver of soil processes in a Mediterranean decomposer system exposed to two moisture levels

The role of biodiversity for soil processes remains poorly understood. Existing evidence suggests that functional diversity rather than species richness is relevant for soil functioning. However, the importance of functional diversity has rarely been assessed simultaneously at more than one trophic level, critically limiting the prediction of consequences of biodiversity loss for soil functioning. In a laboratory microcosm experiment, we tested the hypothesis that increasing functional dissimilarity of both litter‐feeding soil fauna and litter mixtures interactively affects the rates of five different soil processes related to litter decomposition. We created trait‐based functional dissimilarity gradients using five assemblages of two detritivore species and five mixtures of two plant litter species commonly found in Mediterranean shrubland ecosystems of southern France. With increasing drought periods predicted for Mediterranean ecosystems in the future, we additionally included two different watering frequencies to evaluate the impact of drought on soil processes and how drought interacts with functional dissimilarity. The different fauna assemblages and litter mixtures showed strong effects on litter mass loss, soil organic carbon and nitrogen leaching, as well as on soil microbial activities. Up to 20% of the variation in response variables was explained by functional dissimilarity, suggesting an ecologically relevant impact of functional diversity on soil process rates. Detritivore functional dissimilarity tended to have stronger effects when combined with increasingly dissimilar litter mixtures, suggesting that trait dissimilarity interacts across trophic levels. Drought affected several soil processes but did not modify the relationships between functional dissimilarity and process rates. Our results indicate that trait diversity of detritivore assemblages and litter mixtures is an important predictor of soil process rates. The common and easily measurable traits used in our study suggest straightforward application across different types of ecosystems and environmental conditions.     

Partitioning the effect of composition and diversity of tree communities on leaf litter decomposition and soil respiration

The decomposition of plant material is an important ecosystem process influencing both carbon cycling and soil nutrient availability. Quantifying how plant diversity affects decomposition is thus crucial for predicting the effect of the global decline in plant diversity on ecosystem functioning. Plant diversity could affect the decomposition process both directly through the diversity of the litter, and/or indirectly through the diversity of the host plant community and its affect on the decomposition environment. Using a biodiversity experiment with trees in which both functional and taxonomic diversity were explicitly manipulated independently, we tested the effects of the functional diversity and identity of the living trees separately and in combination with the functional diversity and identity of the decomposing litter on rates of litter decomposition and soil respiration. Plant traits, predominantly leaf chemical and physical traits, were correlated with both litter decomposition and soil respiration rates. Surface litter decomposition, quantified by mass loss in litterbags, was best explained by abundance‐weighted mean trait values of tree species from which the litter was assembled (functional identity). In contrast, soil respiration, which includes decomposition of dissolved organic carbon and root respiration, was best explained by the variance in trait values of the host trees (functional diversity). This research provides insight into the effect of loss of tree diversity in forests on soil processes. Such understanding is essential to predicting changes in the global carbon budget brought on by biodiversity loss.     

Traits of dominant plant species drive normalized difference vegetation index in grasslands globally

Aim

Theoretical, experimental and observational studies have shown that biodiversity–ecosystem functioning (BEF) relationships are influenced by functional community structure through two mutually non-exclusive mechanisms: (1) the dominance effect (which relates to the traits of the dominant species); and (2) the niche partitioning effect [which relates to functional diversity (FD)]. Although both mechanisms have been studied in plant communities and experiments at small spatial extents, it remains unclear whether evidence from small-extent case studies translates into a generalizable macroecological pattern. Here, we evaluate dominance and niche partitioning effects simultaneously in grassland systems world-wide.

Location

Two thousand nine hundred and forty-one grassland plots globally.

Time period

2000–2014.

Major taxa studied

Vascular plants.

Methods

We obtained plot-based data on functional community structure from the global vegetation plot database “sPlot”, which combines species composition with plant trait data from the “TRY” database. We used data on the community-weighted mean (CWM) and FD for 18 ecologically relevant plant traits. As an indicator of primary productivity, we extracted the satellite-derived normalized difference vegetation index (NDVI) from MODIS. Using generalized additive models and deviation partitioning, we estimated the contributions of trait CWM and FD to the variation in annual maximum NDVI, while controlling for climatic variables and spatial structure.

Results

Grassland communities dominated by relatively tall species with acquisitive traits had higher NDVI values, suggesting the prevalence of dominance effects for BEF relationships. We found no support for niche partitioning for the functional traits analysed, because NDVI remained unaffected by FD. Most of the predictive power of traits was shared by climatic predictors and spatial coordinates. This highlights the importance of community assembly processes for BEF relationships in natural communities.

Main conclusions

Our analysis provides empirical evidence that plant functional community structure and global patterns in primary productivity are linked through the resource economics and size traits of the dominant species. This is an important test of the hypotheses underlying BEF relationships at the global scale.     

Tree litter functional diversity and nitrogen concentration enhance litter decomposition via changes in earthworm communities

Biodiversity is a major driver of numerous ecosystem functions. However, consequences of changes in forest biodiversity remain difficult to predict because of limited knowledge about how tree diversity influences ecosystem functions. Litter decomposition is a key process affecting nutrient cycling, productivity, and carbon storage and can be influenced by plant biodiversity. Leaf litter species composition, environmental conditions, and the detritivore community are main components of the decomposition process, but their complex interactions are poorly understood. In this study, we tested the effect of tree functional diversity (FD) on litter decomposition in a field experiment manipulating tree diversity and partitioned the effects of litter physiochemical diversity and the detritivore community. We used litterbags with different mesh sizes to separate the effects of microorganisms and microfauna, mesofauna, and macrofauna and monitored soil fauna using pitfall traps and earthworm extractions. We hypothesized that higher tree litter FD accelerates litter decomposition due to the availability of complementary food components and higher activity of detritivores. Although we did not find direct effects of tree FD on litter decomposition, we identified key litter traits and macrodetritivores that explained part of the process. Litter mass loss was found to decrease with an increase in leaf litter carbon:nitrogen ratio. Moreover, litter mass loss increased with an increasing density of epigeic earthworms, with most pronounced effects in litterbags with a smaller mesh size, indicating indirect effects. Higher litter FD and litter nutrient content were found to increase the density of surface‐dwelling macrofauna and epigeic earthworm biomass. Based on structural equation modeling, we conclude that tree FD has a weak positive effect on soil surface litter decomposition by increasing the density of epigeic earthworms and that litter nitrogen‐related traits play a central role in tree composition effects on soil fauna and decomposition.     

Assessing the functional turnover of species assemblages with tailored dissimilarity matrices

It is often suggested that community functional diversity is an appropriate predictive measure of ecosystem functioning, particularly if relevant species traits for the ecological property of interest are carefully selected. However, methods for selecting traits are often based on expert knowledge or on theoretical models of ecosystem functioning, but usually do not include explicitly developed quantitative procedures. Here we propose to construct a so‐called ‘tailored dissimilarity matrix’ between species assemblages to emphasize their functional turnover in response to some user‐defined ecological property. First, a subset of community weighted mean trait values (CWM) is selected by stepwise regression on the ecological process of interest. The selected CWM values are then replaced by the residuals of the least‐squares regressions of each single CWM on the ecological process of interest and pairwise Euclidean distances between the residual values at each sampling site are calculated. We illustrate the advantages of the tailored approach using two distinct plant and bee communities under contrasting fire regimes in temperate forests of southern Switzerland. Our results demonstrated that, unlike for the original CWM values, the tailored approach optimized the degree of functional differentiation among bee and plant species assemblages, i.e. the species functional turnover, with respect to different fire regimes.     

Litter and soil biodiversity jointly drive ecosystem functions

The decomposition of litter and the supply of nutrients into and from the soil are two fundamental processes through which the above- and belowground world interact. Microbial biodiversity, and especially that of decomposers, plays a key role in these processes by helping litter decomposition. Yet the relative contribution of litter diversity and soil biodiversity in supporting multiple ecosystem services remains virtually unknown. Here we conducted a mesocosm experiment where leaf litter and soil biodiversity were manipulated to investigate their influence on plant productivity, litter decomposition, soil respiration, and enzymatic activity in the littersphere. We showed that both leaf litter diversity and soil microbial diversity (richness and community composition) independently contributed to explain multiple ecosystem functions. Fungal saprobes community composition was especially important for supporting ecosystem multifunctionality (EMF), plant production, litter decomposition, and activity of soil phosphatase when compared with bacteria or other fungal functional groups and litter species richness. Moreover, leaf litter diversity and soil microbial diversity exerted previously undescribed and significantly interactive effects on EMF and multiple individual ecosystem functions, such as litter decomposition and plant production. Together, our work provides experimental evidence supporting the independent and interactive roles of litter and belowground soil biodiversity to maintain ecosystem functions and multiple services.     

Shifts in plant functional community composition under hydrological stress strongly decelerate litter decomposition

Litter decomposition is a key process of nutrient and carbon cycling in terrestrial ecosystems. The decomposition process will likely be altered under ongoing climate change, both through direct effects on decomposer activity and through indirect effects caused by changes in litter quality. We studied how hydrological change indirectly affects decomposition via plant functional community restructuring caused by changes in plant species’ relative abundances (community‐weighted mean (CWM) traits and functional diversity). We further assessed how those indirect litter quality effects compare to direct effects. We set up a mesocosm experiment, in which sown grassland communities and natural turf pieces were subjected to different hydrological conditions (dryness and waterlogging) for two growing seasons. Species‐level mean traits were obtained from trait databases and combined with species’ relative abundances to assess functional community restructuring. We studied decomposition of mixed litter from these communities in a common “litterbed.” These indirect effects were compared to effects of different hydrological conditions on soil respiration and on decomposition of standard litter (direct effects). Dryness reduced biomass production in sown communities and natural turf pieces, while waterlogging only reduced biomass in sown communities. Hydrological stress caused profound shifts in species’ abundances and consequently in plant functional community composition. Hydrologically stressed communities had higher CMW leaf dry matter content, lower CMW leaf nitrogen content, and lower functional diversity. Lower CWM leaf N content and functional diversity were strongly related to slower decomposition. These indirect effects paralleled direct effects, but were larger and longer‐lasting. Species mean traits from trait databases had therefore considerable predictive power for decomposition. Our results show that stressful soil moisture conditions, that are likely to occur more frequently in the future, quickly shift species’ abundances. The resulting functional community restructuring will decelerate decomposition under hydrological stress.     

Early subtropical forest growth is driven by community mean trait values and functional diversity rather than the abiotic environment

While functional diversity (FD) has been shown to be positively related to a number of ecosystem functions including biomass production, it may have a much less pronounced effect than that of environmental factors or species‐specific properties. Leaf and wood traits can be considered particularly relevant to tree growth, as they reflect a trade‐off between resources invested into growth and persistence. Our study focussed on the degree to which early forest growth was driven by FD, the environment (11 variables characterizing abiotic habitat conditions), and community‐weighted mean (CWM) values of species traits in the context of a large‐scale tree diversity experiment (BEF‐China). Growth rates of trees with respect to crown diameter were aggregated across 231 plots (hosting between one and 23 tree species) and related to environmental variables, FD, and CWM, the latter two of which were based on 41 plant functional traits. The effects of each of the three predictor groups were analyzed separately by mixed model optimization and jointly by variance partitioning. Numerous single traits predicted plot‐level tree growth, both in the models based on CWMs and FD, but none of the environmental variables was able to predict tree growth. In the best models, environment and FD explained only 4 and 31% of variation in crown growth rates, respectively, while CWM trait values explained 42%. In total, the best models accounted for 51% of crown growth. The marginal role of the selected environmental variables was unexpected, given the high topographic heterogeneity and large size of the experiment, as was the significant impact of FD, demonstrating that positive diversity effects already occur during the early stages in tree plantations.     

Landscape moderation of biodiversity patterns and processes - eight hypotheses

Understanding how landscape characteristics affect biodiversity patterns and ecological processes at local and landscape scales is critical for mitigating effects of global environmental change. In this review, we use knowledge gained from human-modified landscapes to suggest eight hypotheses, which we hope will encourage more systematic research on the role of landscape composition and configuration in determining the structure of ecological communities, ecosystem functioning and services. We organize the eight hypotheses under four overarching themes. Section A: 'landscape moderation of biodiversity patterns' includes (1) the landscape species pool hypothesis-the size of the landscape-wide species pool moderates local (alpha) biodiversity, and (2) the dominance of beta diversity hypothesis-landscape-moderated dissimilarity of local communities determines landscape-wide biodiversity and overrides negative local effects of habitat fragmentation on biodiversity. Section B: 'landscape moderation of population dynamics' includes (3) the cross-habitat spillover hypothesis-landscape-moderated spillover of energy, resources and organisms across habitats, including between managed and natural ecosystems, influences landscape-wide community structure and associated processes and (4) the landscape-moderated concentration and dilution hypothesis-spatial and temporal changes in landscape composition can cause transient concentration or dilution of populations with functional consequences. Section C: 'landscape moderation of functional trait selection' includes (5) the landscape-moderated functional trait selection hypothesis-landscape moderation of species trait selection shapes the functional role and trajectory of community assembly, and (6) the landscape-moderated insurance hypothesis-landscape complexity provides spatial and temporal insurance, i.e. high resilience and stability of ecological processes in changing environments. Section D: 'landscape constraints on conservation management' includes (7) the intermediate landscape-complexity hypothesis-landscape-moderated effectiveness of local conservation management is highest in structurally simple, rather than in cleared (i.e. extremely simplified) or in complex landscapes, and (8) the landscape-moderated biodiversity versus ecosystem service management hypothesis-landscape-moderated biodiversity conservation to optimize functional diversity and related ecosystem services will not protect endangered species. Shifting our research focus from local to landscape-moderated effects on biodiversity will be critical to developing solutions for future biodiversity and ecosystem service management.     

Differential effects of functional traits on aboveground biomass in semi-natural grasslands

Despite increasing evidence on the importance of species functional characteristics for ecosystem processes, two major hypotheses suggest different mechanisms: the ‘mass ratio hypothesis’ assumes that functional traits of the dominant species determine ecosystem processes, while the ‘complementarity hypothesis’ predicts that resource niches may be used more completely when a community is functionally more diverse. Here, we present a method which uses two different groups of biotic predictor variables being (1) abundance‐weighted mean (=aggregated) trait values and (2) functional trait diversity based on Rao's quadratic diversity (FD_Q) to test the competing hypotheses on biodiversity–ecosystem functioning relationships after accounting for co‐varying abiotic factors. We applied this method to data recorded on biodiversity–biomass relationships and environmental variables in 35 semi‐natural temperate grasslands and used a literature‐based matrix of fourteen plant functional traits to assess the explanatory power of models including different sets of predictor variables. Aboveground community biomass did not correlate with species richness. Abiotic factors, in particular soil nitrogen concentration, explained about 50% of variability in aboveground biomass. The best model incorporating functional trait diversity explained only about 30%, while the best model based on aggregated trait values explained about 54% of variability in aboveground biomass. The inclusion of all predictor variable groups in a combined model increased the predictive power to about 75%. This model comprised soil nitrogen concentration as abiotic factor, aggregated traits being indicative for species competitive dominance (rooting depth, leaf distribution, specific leaf area, perennial life cycle) and functional trait diversity in vegetative plant height, leaf area and life cycle. Our study strongly suggests that abiotic factors, trait values of the dominant species and functional trait diversity in combination may best explain differences in aboveground community biomass in natural ecosystems and that their isolated consideration may be misleading.