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Clonal growth of plants is mainly a result of the vegetative growth of organs hidden beneath the soil surface and producing potentially independent vegetative offspring. Clonal traits are difficult to measure due to inaccessibility of the space they inhabit and their morphological diversity. This causes great difficulties with descriptions, standardization of measurements across plant growth forms and, probably, a lack of appropriate questions that should be answered using them. The freely available CLO-PLA database (http://clopla.butbn.cas.cz/) can help to assess the roles of vegetative means of regeneration and spread in plant communities under the effect of various biotic and abiotic filters. It can serve as a source of reference on persistence traits of European temperate flora and, eventually, as a guide for trait sampling in other regions of the world.

Global environmental change affects the sustained provision of a wide set of ecosystem services. Although the delivery of ecosystem services is strongly affected by abiotic drivers and direct land use effects, it is also modulated by the functional diversity of biological communities (the value, range, and relative abundance of functional traits in a given ecosystem). The focus of this article is on integrating the different possible mechanisms by which functional diversity affects ecosystem properties that are directly relevant to ecosystem services. We propose a systematic way for progressing in understanding how land cover change affects these ecosystem properties through functional diversity modifications. Models on links between ecosystem properties and the local mean, range, and distribution of plant trait values are numerous, but they have been scattered in the literature, with varying degrees of empirical support and varying functional diversity components analyzed. Here we articulate these different components in a single conceptual and methodological framework that allows testing them in combination. We illustrate our approach with examples from the literature and apply the proposed framework to a grassland system in the central French Alps in which functional diversity, by responding to land use change, alters the provision of ecosystem services important to local stakeholders. We claim that our framework contributes to opening a new area of research at the interface of land change science and fundamental ecology.

Understanding the influence of the environment on the functional structure of ecological communities is essential to predict the response of biodiversity to global change drivers. Ecological theory suggests that multiple environmental factors shape local species assemblages by progressively filtering species from the regional species pool to local communities. These successive filters should influence the various components of community functional structure in different ways. In this paper, we tested the relative influence of multiple environmental filters on various metrics of plant functional trait structure (i.e. ‘community weighted mean trait’ and components of functional trait diversity, i.e. functional richness, evenness and divergence) in 82 vegetation plots in the Guisane Valley, French Alps. For the 211 sampled species we measured traits known to capture key aspects of ecological strategies amongst vascular plant species, i.e. leaf traits, plant height and seed mass (LHS). A comprehensive information theory framework, together with null model based resampling techniques, was used to test the various environmental effects. Particular community components of functional structure responded differently to various environmental gradients, especially concerning the spatial scale at which the environmental factors seem to operate. Environmental factors acting at a large spatial scale (e.g. temperature) were found to predominantly shape community weighted mean trait values, while fine-scale factors (topography and soil characteristics) mostly influenced functional diversity and the distribution of trait values among the dominant species. Our results emphasize the hierarchical nature of ecological forces shaping local species assemblage: large-scale environmental filters having a primary effect, i.e. selecting the pool of species adapted to a site, and then filters at finer scales determining species abundances and local species coexistence. This suggests that different components of functional community structure will respond differently to environmental change, so that predicting plant community responses will require a hierarchical multi-facet approach.

A methodology for partitioning of biodiversity into α, β and γ components has long been debated, resulting in different mathematical frameworks. Recently, use of the Rao quadratic entropy index has been advocated since it allows comparison of various facets of diversity (e.g. taxonomic, phylogenetic and functional) within the same mathematical framework. However, if not well implemented, the Rao index can easily yield biologically meaningless results and lead into a mathematical labyrinth. As a practical guideline for ecologists, we present a critical synthesis of diverging implementations of the index in the recent literature and a new extension of the index for measuring β-diversity. First, we detail correct computation of the index that needs to be applied in order not to obtain negative β-diversity values, which are ecologically unacceptable, and elucidate the main approaches to calculate the Rao quadratic entropy at different spatial scales. Then, we emphasize that, similar to other entropy measures, the Rao index often produces lower-than-expected β-diversity values. To solve this, we extend a correction based on equivalent numbers, as proposed by Jost (2007), to the Rao index. We further show that this correction can be applied to additive partitioning of diversity and not only its multiplicative form. These developments around the Rao index open up an exciting avenue to develop an estimator of turnover diversity across different environmental and temporal scales, allowing meaningful comparisons of partitioning across species, phylogenetic and functional diversities within the same mathematical framework. We also propose a set of R functions, based on existing developments, which perform different key computations to apply this framework in biodiversity science.

1. The mechanisms driving nonrandom assembly patterns in plant communities have long been of interest in ecological research. Competing ecological theories predict that coexisting species may either be more functionally dissimilar than expected by chance (with functional 'divergence' mainly reflecting niche differentiation) or be functionally more similar than expected (with functional 'convergence' reflecting either the outcome of environmental filtering or weaker-competitor exclusion effects). Assembly patterns are usually assessed at a single scale and disturbance regime, whereas considering different spatial scales and disturbance regimes may clarify the underlying assembly mechanisms. 2. We tested the prediction that convergence and divergence are scale-and disturbance-dependent in grazed and abandoned species-rich dry grasslands within a 22 km² landscape in south-eastern Sweden. Convergence and divergence were tested for plant species' traits and phylogenetic relationships at three nested spatial scales: within 412 plots (50 × 50 cm, divided into 10 × 10 cm subplots), within 117 grassland patches (from 0.02 to 11.63 ha) and within the whole landscape (across patches). 3. At the finest scale (10 × 10 cm subplots within plots), coexisting species were more different than expected by chance (divergence), both functionally and phylogenetically, suggesting niche differentiation. At the intermediate scale (50 × 50 cm plots within patches), coexisting species showed convergence, suggesting environmental filtering. No significant deviations from random expectations were detected at the broadest scale (patches within the 22 km² landscape) -suggesting the prevalence of dispersal limitation at this scale. The fact that nonrandom patterns were particularly evident under grazed conditions is consistent with the prediction that assembly patterns are disturbance dependent. 4. Synthesis. This study shows that multiple trait-based assembly processes operate simultaneously in species-rich communities, across spatial scales and disturbance regimes. The results support earlier theoretical predictions that divergence between coexisting species may be an important driver of community assembly, particularly at finer spatial scales, where species compete for the same local resources. In contrast, environmental filtering is expected at broader spatial scales, where species growing in particular environmental conditions share traits that are adaptive under those conditions. Within given habitat types, dispersal limitation may, however, override environmental filtering at increasing spatial scales of observation.

FdB, WT and SL were partially supported by the DIVERSITALP project (ANR 07 BDIV 014). The research was funded by the project DIVHERBE from the French ACI-ECOGER programme, the Czech projects LC 06073, GACR(206/06/ 0098) and MSMT6007665801, the EU projects RUBICODE (FP6, No. 036890), PASTUS-INTERREG (I3A-4-147-E, INTERREG III-A programme, EU) and EcoChange (Challenges in assessing and forecasting biodiversity and ecosystem changes in Europe, No: GOCE-CT-2003-506675).

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.

The promise of “trait-based” plant ecology is one of generalized prediction across organizational and spatial scales, independent of taxonomy. This promise is a major reason for the increased popularity of this approach. Here, we argue that some important foundational assumptions of trait-based ecology have not received sufficient empirical evaluation. We identify three such assumptions and, where possible, suggest methods of improvement: (i) traits are functional to the degree that they determine individual fitness, (ii) intraspecific variation in functional traits can be largely ignored, and (iii) functional traits show general predictive relationships to measurable environmental gradients.

The loss of biodiversity is thought to have adverse effects on multiple ecosystem functions, including the decline of community stability. Decreased diversity reduces the strength of the portfolio effect, a mechanism stabilizing community temporal fluctuations. Community stability is also expected to decrease with greater variability in individual species populations and with synchrony of their fluctuations. In semi-natural meadows, eutrophication is one of the most important drivers of diversity decline; it is expected to increase species fluctuations and synchrony among them, all effects leading to lower community stability. With a 16-year time series of biomass data from a temperate species-rich meadow with fertilization and removal of the dominant species, we assessed population biomass temporal (co)variation under different management types and competition intensity, and in relation to species functional traits and to species diversity. Whereas the effect of dominant removal was relatively small (with a tendency toward lower stability), fertilization markedly decreased community stability (i.e., increased coefficient of variation in the total biomass) and species diversity. On average, the fluctuations of individual populations were mutually independent, with a slight tendency toward synchrony in unfertilized plots, and a tendency toward compensatory dynamics in fertilized plots and no effects of removal. The marked decrease of synchrony with fertilization, contrary to the majority of the results reported previously, follows the predictions of increased compensatory dynamics with increased asymmetric competition for light in a more productive environment. Synchrony increased also with species functional similarity stressing the importance of shared ecological strategies in driving similar species responses to weather fluctuations. As expected, the decrease of temporal stability of total biomass was mainly related to the decrease of species richness, with its effect remaining significant also after accounting for fertilization. The weakening of the portfolio effect with species richness decline is a crucial driver of community destabilization. However, the positive effect of species richness on temporal stability of total biomass was not due to increased compensatory dynamics, since synchrony increased with species richness. This shows that the negative effect of eutrophication on community stability does not operate through increasing synchrony, but through the reduction of diversity.

Question: Which functional diversity indices have the power to reveal changes in community assembly processes along abiotic stress gradients? Is their power affected by stochastic processes and variations in species richness along stress gradients? Methods: We used a simple community assembly model to explore the power of functional diversity indices across a wide range of ecological contexts. The model assumes that with declining stress the influence of niche complementarity on species fitness increases while that of environmental filtering decreases. We separately incorporated two trait-independent stochastic processes — mass and priority effects — in simulating species occurrences and abundances along a hypothetical stress gradient. We ran simulations where species richness was constant along the gradient, or increased, decreased or varied randomly with declining stress. We compared observed values for two indices of functional richness — total functional dendrogram length (FD) and convex hull volume (FRic) — with a matrix-swap null model (yielding indices SESFD and SESFRic) to remove any trivial effects of species richness. We also compared two indices that measure both functional richness and functional divergence — Rao quadratic entropy (Rao) and functional dispersion (FDis) — with a null model that randomizes abundances across species but within communities. This converts them to pure measures of functional divergence (SESRao and SESFDis). Results: When mass effects operated, only SESRao and SESFDis gave reasonable power, irrespective of how species richness varied along the stress gradient. FD, FRic, Rao and FDis had low power when species richness was constant, and variation in species richness greatly influenced their power. SESFRic and SESFD were unaffected by variation in species richness. When priority effects operated, FRic, SESFRic, Rao and FDis had good power and were unaffected by variation in species richness. Variation in species richness greatly affected FD and SESFD. SESRao and SESFDis had low power in the priority effects model but were unaffected by variation in species richness. Conclusions: Our results demonstrate that a reliable test for changes in assembly processes along stress gradients requires functional diversity indices measuring either functional richness or functional divergence. We recommend using SESFRic as a measure of functional richness and either SESRao or SESFDis (which are very closely related mathematically) as a measure of functional divergence. Used together, these indices of functional richness and functional divergence provide good power to test for increasing niche complementarity with declining stress across a broad range of ecological contexts.