Functional diversity changes during tropical forest succession

Functional diversity (FD) ‘those components of biodiversity that influence how an ecosystem operates or functions’ is a promising tool to assess the effect of biodiversity loss on ecosystem functioning. FD has received ample theoretical attention, but empirical studies are limited. We evaluate changes in species richness and FD during tropical secondary forest succession after shifting cultivation in Mexico. We also test whether species richness is a good predictor of FD. FD was calculated based on a combination of nine functional traits, and based on two individual traits important for primary production (specific leaf area) and carbon sequestration (wood density). Stand basal area was a good predictor of successional changes in diversity and FD, in contrast to fallow age. Incidence-based FD indices increased logarithmically with stand basal area, but FD weighted by species’ importance values lacked pattern with succession. Species richness and diversity are strong predictors of FD when all traits were considered; linear relationships indicate that all species are equally functionally complementary, suggesting there is little functional redundancy. In contrast, when FD was calculated for individual traits and weighted for abundances, species richness may underestimate FD.Selection of functional trait(s) critically determines FD, with large consequences for studies relating biodiversity to ecosystem functioning. Careful consideration of the traits required to capture the ecosystem process of interest is thus essential.     

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.     

Testing the Link between Functional Diversity and Ecosystem Functioning in a Minnesota Grassland Experiment

The functional diversity of a community can influence ecosystem functioning and reflects assembly processes. The large number of disparate metrics used to quantify functional diversity reflects the range of attributes underlying this concept, generally summarized as functional richness, functional evenness, and functional divergence. However, in practice, we know very little about which attributes drive which ecosystem functions, due to a lack of field-based tests. Here we test the association between eight leading functional diversity metrics (Rao’s Q, FD, FDis, FEve, FDiv, convex hull volume, and species and functional group richness) that emphasize different attributes of functional diversity, plus 11 extensions of these existing metrics that incorporate heterogeneous species abundances and trait variation. We assess the relationships among these metrics and compare their performances for predicting three key ecosystem functions (above- and belowground biomass and light capture) within a long-term grassland biodiversity experiment. Many metrics were highly correlated, although unique information was captured in FEve, FDiv, and dendrogram-based measures (FD) that were adjusted by abundance. FD adjusted by abundance outperformed all other metrics in predicting both above- and belowground biomass, although several others also performed well (e.g. Rao’s Q, FDis, FDiv). More generally, trait-based richness metrics and hybrid metrics incorporating multiple diversity attributes outperformed evenness metrics and single-attribute metrics, results that were not changed when combinations of metrics were explored. For light capture, species richness alone was the best predictor, suggesting that traits for canopy architecture would be necessary to improve predictions. Our study provides a comprehensive test linking different attributes of functional diversity with ecosystem function for a grassland system.     

Functional identity is the main driver of diversity effects in young tree communities

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.     

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.     

Loss of functional diversity under land use intensification across multiple taxa

Land use intensification can greatly reduce species richness and ecosystem functioning. However, species richness determines ecosystem functioning through the diversity and values of traits of species present. Here, we analyze changes in species richness and functional diversity (FD) at varying agricultural land use intensity levels. We test hypotheses of FD responses to land use intensification in plant, bird, and mammal communities using trait data compiled for 1600+ species. To isolate changes in FD from changes in species richness we compare the FD of communities to the null expectations of FD values. In over one-quarter of the bird and mammal communities impacted by agriculture, declines in FD were steeper than predicted by species number. In plant communities, changes in FD were indistinguishable from changes in species richness. Land use intensification can reduce the functional diversity of animal communities beyond changes in species richness alone, potentially imperiling provisioning of ecosystem services.     

植物群落功能多样性计算方法

以性状为基础的功能多样性指数在预测生态系统功能中起到越来越重要的作用.本文对近年来陆续出现的植物群落功能多样性指数进行综述.依据物种多样性指数的组成,功能多样性指数分为功能丰富度、功能均匀度和功能离散度指数.介绍了这3类指数的计算方法,有助于更好、更准确地理解“生物多样性-环境-生态系统功能”的关系.     

Impact of land-use intensity on the conservation of functional and phylogenetic diversity in temperate semi-natural plant communities

The earth is facing a worldwide decline in biodiversity, with land-use change identified as one of the most important drivers. There is evidence that the loss of diversity has a significant impact on ecosystem functioning. Earlier research focused on species richness, but more recent, functional and phylogenetic diversity came into the picture as the stronger determinants of ecosystem processes. The effects of increasing land-use intensity on functional (FD) and phylogenetic diversity (PD), however, are still poorly understood. We studied how FD and PD are affected by land-use intensity in temperate plant communities. Our results show that land-use intensity has a clear impact on species richness, but also affects functional and phylogenetic diversity. Intensive agricultural areas fail to support high and sustainable levels of functional and phylogenetic diversity. These results highlight the need for the protection of biodiversity in nature reserves and the conservation of areas with extensive agricultural practices. Because species richness may influence the measures of functional and phylogenetic diversity, we compared the observed FD and PD values with random values generated with a matrix-swap null model. The observed discrepancy between species loss and the loss of FD and PD calls for an integrated approach to biodiversity conservation, in which the different components of biodiversity are considered together.     

Functional diversity predicts overyielding effect of species combination on primary productivity

The effect of biodiversity on ecosystem functioning has proven variable both within and among manipulative studies. Species richness is the most commonly used measure of biodiversity in such studies, but the range of species’ functional traits (functional diversity), not the number of species per se, likely underpins a key mechanistic link between species richness and ecosystem functioning. However, the majority of experiments that have examined the effect of functional diversity have manipulated functional group richness, an approach recognised to suffer numerous limitations. Continuous measures of functional diversity avoid many of these limitations, but the relationship between continuous functional diversity and the magnitude of ecosystem processes has been largely untested. Using one vs two‐species mixtures of rock pool macroalgae as a model, we conducted a field experiment to determine the effect of a continuous measure of functional diversity (functional attribute diversity, FAD, the degree of functional differentiation based on four functional traits) on the magnitude of net primary productivity and overyielding, based upon two alternative null‐models. The total magnitude of productivity was largely determined by the identity of species present, not FAD. However, FAD proved to be a good predictor of overyielding (variation in productivity after the dominant effects of species identity had been accounted for). Furthermore, despite differences in the mean magnitude of the effect of combining species, the positive relationship between FAD and overyielding was consistent according to both additive and substitutive null‐models. Our findings imply that whilst knowledge of species’ independent contributions remains indispensable in the prediction of biotic effects on ecosystem functioning within a trophic level, continuous measures of functional diversity should be used as a supplementary tool to predict the magnitude of overyielding, thereby refining predictions.     

Relationships between functional diversity and ecosystem functioning: A review

Functional diversity, which is the value, variation and distribution of traits in a community assembly, is an important component of biodiversity. Functional diversity is generally viewed as a key to understand ecosystem and community functioning. There are three components of functional diversity, i.e. functional richness, evenness and divergence. Functional diversity and species diversity can be either positively or negatively correlated, or uncorrelated, depending on the environmental conditions and disturbance intensity. Ecosystem functioning includes ecosystem processes, ecosystem properties and ecosystem stability. The diversity hypothesis and the mass ratio hypothesis are the two major hypotheses of explaining the effect of functional diversity on ecosystem functioning, diversity hypothesis reflects that organisms and their functional traits in a assemblage effect on ecosystem functioning by the complementarity of using resources, and mass ratio hypothesis emphasises the identify of the dominant species in a assemblage. These two hypotheses do not contradict each other and instead they reflect the two different sides of functional diversity and functional composition. The effect of functional diversity on ecosystem functioning also depends on abiotic factors, perturbation, management actions, etc. Function diversity potentially influences ecosystem service and management by effecting on ecosystem functioning. Ecosystem management groups should include functional diversity in their scheme and not just species richness.     

Ecosystem Functions across Trophic Levels Are Linked to Functional and Phylogenetic Diversity

In experimental systems, it has been shown that biodiversity indices based on traits or phylogeny can outperform species richness as predictors of plant ecosystem function. However, it is unclear whether this pattern extends to the function of food webs in natural ecosystems. Here we tested whether zooplankton functional and phylogenetic diversity explains the functioning of 23 natural pond communities. We used two measures of ecosystem function: (1) zooplankton community biomass and (2) phytoplankton abundance (Chl a). We tested for diversity-ecosystem function relationships within and across trophic levels. We found a strong correlation between zooplankton diversity and ecosystem function, whereas local environmental conditions were less important. Further, the positive diversity-ecosystem function relationships were more pronounced for measures of functional and phylogenetic diversity than for species richness. Zooplankton and phytoplankton biomass were best predicted by different indices, suggesting that the two functions are dependent upon different aspects of diversity. Zooplankton community biomass was best predicted by zooplankton trait-based functional richness, while phytoplankton abundance was best predicted by zooplankton phylogenetic diversity. Our results suggest that the positive relationship between diversity and ecosystem function can extend across trophic levels in natural environments, and that greater insight into variation in ecosystem function can be gained by combining functional and phylogenetic diversity measures.     

Plant diversity induces shifts in the functional structure and diversity across trophic levels

Changes to primary producer diversity can cascade up to consumers and affect ecosystem processes. Although the effect of producer diversity on higher trophic groups have been studied, these studies often quantify taxonomy‐based measures of biodiversity, like species richness, which do not necessarily reflect the functioning of these communities. In this study, we assess how plant species richness affects the functional composition and diversity of higher trophic levels and discuss how this might affect ecosystem processes, such as herbivory, predation and decomposition. Based on six different consumer traits, we examined the functional composition of arthropod communities sampled in experimental plots that differed in plant species richness. The two components we focused on were functional variation in the consumer community structure (functional structure) and functional diversity, expressed as functional richness, evenness and divergence. We found a consistent positive effect of plant species richness on the functional richness of herbivores, carnivores, and omnivores, but not decomposers, and contrasting patterns for functional evenness and divergence. Increasing plant species richness shifted the omnivore community to more predatory and less mobile species, and the herbivore community to more specialized and smaller species. This was accompanied by a shift towards more species occurring in the vegetation than in the ground layer. Our study shows that plant species richness strongly affects the functional structure and diversity of aboveground arthropod communities. The observed shifts in body size (herbivores), specialization (herbivores), and feeding mode (omnivores) together with changes in the functional diversity may underlie previously observed increases in herbivory and predation in plant communities of higher diversity.     

Functional richness outperforms taxonomic richness in predicting ecosystem functioning in natural phytoplankton communities

相似文献   

生态学的多样性指数:功能与系统发育

生物多样性是生态学的核心问题。传统的多样性指数仅包含物种数和相对多度的信息,这类基于分类学的多样性指数并不能很好地帮助理解群落构建和生态系统功能。不同物种对群落构建和生态系统功能所起到的作用类型和贡献也不完全相同,且物种在生态过程中的作用和贡献往往与性状密切相关,因此功能多样性已经成为反映物种群落构建、干扰以及环境因素对群落影响的重要指标。同时,由于亲缘关系相近的物种往往具有相似的性状,系统发育多样性也可以作为功能多样性的一个替代。功能多样性和系统发育多样性各自具有优缺点,但二者均比分类多样性更能揭示群落和生态系统的构建、维持与功能。     

Biodiversity effects on ecosystem functioning in a 15-year grassland experiment: Patterns,mechanisms, and open questions

In the past two decades, a large number of studies have investigated the relationship between biodiversity and ecosystem functioning, most of which focussed on a limited set of ecosystem variables. The Jena Experiment was set up in 2002 to investigate the effects of plant diversity on element cycling and trophic interactions, using a multi-disciplinary approach. Here, we review the results of 15 years of research in the Jena Experiment, focussing on the effects of manipulating plant species richness and plant functional richness. With more than 85,000 measures taken from the plant diversity plots, the Jena Experiment has allowed answering fundamental questions important for functional biodiversity research.First, the question was how general the effect of plant species richness is, regarding the many different processes that take place in an ecosystem. About 45% of different types of ecosystem processes measured in the ‘main experiment’, where plant species richness ranged from 1 to 60 species, were significantly affected by plant species richness, providing strong support for the view that biodiversity is a significant driver of ecosystem functioning. Many measures were not saturating at the 60-species level, but increased linearly with the logarithm of species richness. There was, however, great variability in the strength of response among different processes. One striking pattern was that many processes, in particular belowground processes, took several years to respond to the manipulation of plant species richness, showing that biodiversity experiments have to be long-term, to distinguish trends from transitory patterns. In addition, the results from the Jena Experiment provide further evidence that diversity begets stability, for example stability against invasion of plant species, but unexpectedly some results also suggested the opposite, e.g. when plant communities experience severe perturbations or elevated resource availability. This highlights the need to revisit diversity–stability theory.Second, we explored whether individual plant species or individual plant functional groups, or biodiversity itself is more important for ecosystem functioning, in particular biomass production. We found strong effects of individual species and plant functional groups on biomass production, yet these effects mostly occurred in addition to, but not instead of, effects of plant species richness.Third, the Jena Experiment assessed the effect of diversity on multitrophic interactions. The diversity of most organisms responded positively to increases in plant species richness, and the effect was stronger for above- than for belowground organisms, and stronger for herbivores than for carnivores or detritivores. Thus, diversity begets diversity. In addition, the effect on organismic diversity was stronger than the effect on species abundances.Fourth, the Jena Experiment aimed to assess the effect of diversity on N, P and C cycling and the water balance of the plots, separating between element input into the ecosystem, element turnover, element stocks, and output from the ecosystem. While inputs were generally less affected by plant species richness, measures of element stocks, turnover and output were often positively affected by plant diversity, e.g. carbon storage strongly increased with increasing plant species richness. Variables of the N cycle responded less strongly to plant species richness than variables of the C cycle.Fifth, plant traits are often used to unravel mechanisms underlying the biodiversity–ecosystem functioning relationship. In the Jena Experiment, most investigated plant traits, both above- and belowground, were plastic and trait expression depended on plant diversity in a complex way, suggesting limitation to using database traits for linking plant traits to particular functions.Sixth, plant diversity effects on ecosystem processes are often caused by plant diversity effects on species interactions. Analyses in the Jena Experiment including structural equation modelling suggest complex interactions that changed with diversity, e.g. soil carbon storage and greenhouse gas emission were affected by changes in the composition and activity of the belowground microbial community. Manipulation experiments, in which particular organisms, e.g. belowground invertebrates, were excluded from plots in split-plot experiments, supported the important role of the biotic component for element and water fluxes.Seventh, the Jena Experiment aimed to put the results into the context of agricultural practices in managed grasslands. The effect of increasing plant species richness from 1 to 16 species on plant biomass was, in absolute terms, as strong as the effect of a more intensive grassland management, using fertiliser and increasing mowing frequency. Potential bioenergy production from high-diversity plots was similar to that of conventionally used energy crops. These results suggest that diverse ‘High Nature Value Grasslands’ are multifunctional and can deliver a range of ecosystem services including production-related services.A final task was to assess the importance of potential artefacts in biodiversity–ecosystem functioning relationships, caused by the weeding of the plant community to maintain plant species composition. While the effort (in hours) needed to weed a plot was often negatively related to plant species richness, species richness still affected the majority of ecosystem variables. Weeding also did not negatively affect monoculture performance; rather, monocultures deteriorated over time for a number of biological reasons, as shown in plant-soil feedback experiments.To summarize, the Jena Experiment has allowed for a comprehensive analysis of the functional role of biodiversity in an ecosystem. A main challenge for future biodiversity research is to increase our mechanistic understanding of why the magnitude of biodiversity effects differs among processes and contexts. It is likely that there will be no simple answer. For example, among the multitude of mechanisms suggested to underlie the positive plant species richness effect on biomass, some have received limited support in the Jena Experiment, such as vertical root niche partitioning. However, others could not be rejected in targeted analyses. Thus, from the current results in the Jena Experiment, it seems likely that the positive biodiversity effect results from several mechanisms acting simultaneously in more diverse communities, such as reduced pathogen attack, the presence of more plant growth promoting organisms, less seed limitation, and increased trait differences leading to complementarity in resource uptake. Distinguishing between different mechanisms requires careful testing of competing hypotheses. Biodiversity research has matured such that predictive approaches testing particular mechanisms are now possible.     

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 .     

A trait-based experimental approach to understand the mechanisms underlying biodiversity–ecosystem functioning relationships

Plant functional characteristics may drive plant species richness effects on ecosystem processes. Consequently, the focus of biodiversity–ecosystem functioning (BEF) experiments has expanded from the manipulation of plant species richness to manipulating functional trait composition. Involving ecophysiological plant traits in the experimental design might allow for a better understanding of how species loss alters ecosystem processes. Here we provide the theoretical background, design and first results of the ‘Trait-Based Biodiversity Experiment’ (TBE), established in 2010 that directly manipulates the trait composition of experimental plant communities.Analysis of six plant traits related to resource acquisition and use were analyzed using principal component analysis of 60 grassland species. The resulting two main axes describe gradients in functional similarity, and were used as the basis for designing plant communities with different functional and species diversity levels. Using such an approach allowed us to manipulate different levels of complementarity in spatial and temporal plant resource acquisition. In contrast to previous biodiversity experiments, the TBE is designed according to more realistic scenarios of non-random species loss along orthogonal axes of species trait dissimilarities. This allows us to tease apart the relative importance of selection and complementarity effects on multiple ecosystem processes, and to mechanistically study the consequences of plant community simplification.     

Predicting productivity in tropical reservoirs: The roles of phytoplankton taxonomic and functional diversity

Primary productivity is intimately linked with biodiversity and ecosystem functioning. Much of what is known today about such relationship has been based on the manipulation of species richness. Other facets of biodiversity, such as functional diversity, have been neglected within this framework, particularly in freshwater systems. We assess the adequacy of different diversity measures, from species richness and evenness, to functional groups richness and functional diversity indices, to predict primary productivity in 19 tropical reservoirs of central Brazil, built to generate hydroelectric energy. We applied linear mixed models (and model selection based on the Akaike’s information criterion) to achieve our goal, using chlorophyll-a concentration as a surrogate for primary productivity. A total of 412 species were collected in this study. Overall we found a positive relation between productivity and diversity, with functional evenness representing the only exception. The most parsimonious models never included functional group classifications, with at least one continuous measure of functional diversity being present in many models. The best model included only species richness and explained 24.1% of variability in productivity. We therefore advise the use of species richness as an indicator of productivity in tropical freshwater environments. However, since the productivity–diversity relationship is known to be scale dependent, we recommend the use of continuous measures of functional diversity in future biodiversity and ecosystem functioning studies, in order to be certain that all functional differences between communities are being accounted for.     

Functional biodiversity of macroinvertebrate assemblages along major ecological gradients of boreal headwater streams

1. Biodiversity–environment relationships are increasingly well‐understood in the context of species richness and species composition, whereas other aspects of biodiversity, including variability in functional diversity (FD), have received rather little rigorous attention. For streams, most studies to date have examined either taxonomic assemblage patterns or have experimentally addressed the importance of species richness for ecosystem functioning. 2. I examined the relationships of the functional biodiversity of stream macroinvertebrates to major environmental and spatial gradients across 111 boreal headwater streams in Finland. Functional biodiversity encompassed functional richness (FR – the number of functional groups derived from a combination of functional feeding groups and habit trait groups), FD – the number of functional groups and division of individuals among these groups, and functional evenness (FE – the division of individuals among functional groups). Furthermore, functional structure (FS) comprised the composition and abundance of functional groups at each site. 3. FR increased with increasing pH, with additional variation related to moss cover, total nitrogen, water colour and substratum particle size. FD similarly increased with increasing pH and decreased with increasing canopy cover. FE decreased with increasing canopy cover and water colour. Significant variation in FS was attributable to pH, stream width, moss cover, substratum particle size, nitrogen, water colour with the dominant pattern in FS being related to the increase of shredder‐sprawlers and the decrease of scraper‐swimmers in acidic conditions. 4. In regression analysis and redundancy analysis, variation in functional biodiversity was not only related to local environmental factors, but a considerable proportion of variability was also attributable to spatial patterning of environmental variables and pure spatial gradients. For FR, 23.4% was related to pure environmental effects, 15.0% to shared environmental and spatial effects and 8.0% to spatial trends. For FD, 13.8% was attributable to environmental effects, 15.2% to shared environmental and spatial effects and 5% to spatial trends. For FE, 9.0% was related to environmental variables, 12.7% to shared effects of environmental and spatial variables and 4.5% to spatial variables. For FS, 13.5% was related to environmental effects, 16.9% to shared environmental and spatial effects and 15.4% to spatial trends. 5. Given that functional biodiversity should portray variability in ecosystem functioning, one might expect to find functionally rather differing ecosystems at the opposite ends of major environmental gradients (e.g. acidity, stream size). However, the degree to which variation in the functional biodiversity of stream macroinvertebrates truly portrays variability in ecosystem functioning is difficult to judge because species traits, such as feeding roles and habit traits, are themselves strongly affected by the habitat template. 6. If functional characteristics show strong responses to natural environmental gradients, they also are likely to do so to anthropogenic environmental changes, including changes in habitat structure, organic inputs and acidifying elements. However, given the considerable degree of spatial structure in functional biodiversity, one should not expect that only the local environment and anthropogenic changes therein are responsible for this variability. Rather, the spatial context, as well as natural variability along environmental gradients, should also be explicitly considered in applied research.