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Changes in phenology induced by climate change occur across the globe with important implications for ecosystem functioning and services, species performance and trophic interactions. Much of the work on phenology, especially leaf out and flowering, has been conducted on woody plant species. Less is known about the responses in phenology of herbaceous species induced by global change even though they represent a large and important part of biodiversity worldwide. A globally coordinated research effort is needed to understand the drivers and implications of such changes and to predict effects of global change on plant species phenology and related ecosystem processes. Here, we present the rationale of the PhenObs initiative-botanical gardens as a global phenological observation network. The initiative aims to collect data on plant phenology in botanical gardens which will be used alongside information on plant traits and site conditions to answer questions related to the consequences of global change: What is the variation in plant phenology in herbaceous species across the growing season and in response to changes in climate? How can plant phenology be predicted from species' trait composition, provenance, position and extent of the distribution range and species' phylogeny? What are the implications of this variation with respect to species performance and assembly, biotic interactions (e.g. plant-pollinator interactions) as well as ecosystem processes and services under changing land use and climate? Here, we lay out the development of a straightforward protocol that is appropriate for monitoring phenology across a vast diversity of growth forms of herbaceous species from various habitats and geographical regions. To focus on a key number of stages necessary to capture all aspects of plant species phenology, we analysed associations between 14 phenological stages. These data were derived from a 2-year study on 199 species in four German botanical gardens. Based on the relationships of the phenological stages, we propose to monitor three vegetative stages ('initial growth', 'leaves unfolding' and 'senescence') and two reproductive stages ('flowers open' and 'ripe fruits') to fully capture herbaceous species phenology. A free Plain Language Summary can be found within the Supporting Information of this article.

Patterns of flowering phenology, i.e. first and last flowering day (FFD and LFD) and flowering duration (FD) govern plant pollination and reproduction. Most research has focused on FFD whereas LFD, FD and second flowering events were neglected although they are crucial events. To assess whether flowering patterns change species-specifically with changing abiotic conditions and whether these are related to leaf persistence, reproduction and pollination syndrome, we monitored flowering phenology (including also second flowering events) of 29 herbaceous species along two elevational gradients ranging from 700 to 1800 m a.s.l. in two consecutive years. FFD was delayed with increasing elevation. LFD followed two alternative strategies: species which ceased flowering early in the season delayed LFD with increasing elevation, species which ceased flowering late did not change LFD. FD decreased with increasing elevation in most species. The timing of flowering phenology had strong influences on the intensity of change along the elevational gradient and other stages of flowering phenology. Many species showed a second flowering event which occurred irrespective of elevation, suggesting a link to unsuccessful pollination rather than temperature. Life history strategies helped explain patterns of flowering phenology and a species’ ability to track changes in abiotic conditions, e.g. evergreen species started to flower earlier than summer green species and insect-pollinated species were able to expand FD most. These findings give insight into species-specific changes in phenological patterns and thus plant performance with changing environment. They should be considered when assessing the consequences of environmental change at both the community and ecosystem level.

Plant functional traits can predict community assembly and ecosystem functioning and are thus widely used in global models of vegetation dynamics and land–climate feedbacks. Still, we lack a global understanding of how land and climate affect plant traits. A previous global analysis of six traits observed two main axes of variation: (1) size variation at the organ and plant level and (2) leaf economics balancing leaf persistence against plant growth potential. The orthogonality of these two axes suggests they are differently influenced by environmental drivers. We find that these axes persist in a global dataset of 17 traits across more than 20,000 species. We find a dominant joint effect of climate and soil on trait variation. Additional independent climate effects are also observed across most traits, whereas independent soil effects are almost exclusively observed for economics traits. Variation in size traits correlates well with a latitudinal gradient related to water or energy limitation. In contrast, variation in economics traits is better explained by interactions of climate with soil fertility. These findings have the potential to improve our understanding of biodiversity patterns and our predictions of climate change impacts on biogeochemical cycles., Nature Ecology & Evolution, 6 (1), ISSN:2397-334X

The intraspecific plant trait variability (ITV) is key for many ecological processes, but large-scale analysis of co-occurring species are rare. Here we studied ITV of 14 plant traits for five co-occurring species across 56 European dry grassland sites – large parts of the species distribution ranges. We evaluated variation in ITV relative to between species trait variability (BTV) with a particular emphasis on how within versus between population variability contributes to ITV. We performed this analysis trait by trait (univariate) and by considering hypervolumes in the multi-variate trait space. We also tested whether climatic variables can be used to predict between population variation in traits. For the trait by trait analysis, for 9 out of 14 traits ITV exceeded BTV, which suggests an extraordinary large role of ITV. However, when considering all traits jointly the hypervolume occupied by ITV was only 23% of overall trait variability. We found comparatively small effects of within population variability in the uni- as well as in the multivariate analysis decreasing rapidly with increasing number of considered traits. The dominance of between population variability suggests that ITV is mainly driven by environmental effects rather than local biotic interactions and microsite effects. ITV of leaf chemical traits was related to precipitation and growing degree days until sampling ITV can be substantial compared to BTV, in particular when considering single traits, such that ITV should be considered in trait-based research. Since the importance of ITV appears to decrease when considering multiple traits jointly, using species mean trait values is more appropriate for multi-variate trait analysis. Because ITV varied mainly between populations rather than within populations, and was related to climatic conditions, we suggest that ITV could be accounted for in future trait based research by including environmental covariates in hierarchical models of trait variation

Moisture availability is the main limiting factor of plant growth and biomass production in arid and semi-arid grasslands. The question whether plant responses to changing precipitation are species-specific, or change over entire plant communities is still controversial. Our study focussed on intraspecific changes in the plant traits canopy height, plant width, specific leaf area, chlorophyll fluorescence, performance index, and individual biomass of three congeneric species pairs with changing precipitation in Mongolian rangelands, covering a gradient from the desert to the forest steppes. Using this trait data set, we focussed on three questions: (i) Is the replacement of congeneric species along an environmental gradient also reflected in their trait values? (ii) Can intraspecific trait variation patterns be derived from patterns in species abundances, i.e., are trait values optimal where species are most abundant? (iii) Is the within-population trait variability lowest in populations growing under very dry conditions, i.e., under highest environmental stress, caused by stronger filtering? We tested the responses of the six traits to changing precipitation according to species’ identity and abundance. We found unimodal relationships between most of the species’ traits and precipitation, and strong associations between species abundances and trait values, but not for all investigated species. Trait variability did not significantly differ between populations from different positions along the precipitation gradient. Our results highlight that species show multiple or even opposite trait responses along the precipitation gradient. It thus remains challenging to predict how plant distributions will shift under changing environmental conditions based on their trait composition.

Frost resistance (FR) is a highly adaptive trait and important for plant performance, survival and distribution. While overall seasonal changes in the frost resistance of herbaceous species are well documented, knowledge of the variability during the growth period is scarce. Responses could be expected due to differences in temperature yet investment in frost resistance might be at the expense of plant performance. To analyse temporal and spatial (i.e. same date but differing temperatures) variability, FR of leaves of six herbaceous species on five sampling dates were assessed along an elevational gradient in the northern limestone Alps. We used chlorophyll fluorescence techniques to calculate the lethal temperature of 90% of the population (LT90) thereof. To test the association with plant performance, we measured eco-morphological leaf traits (specific leaf area (SLA), leaf dry matter content, leaf phosphorous and magnesium content as well as stomatal pore area index (SPI)) in parallel. We found that FR as well as leaf traits exhibit a strong temporal variation whereas spatial variability was low. When analysing the relationship of FR to leaf traits we found that SLA as a proxy of growth rate was negatively associated with FR indicating a trade-off between growth and resistance, whereas SPI showed a positive relationship to FR. This finding gives further insight into the variability of traits and will help to improve predictions concerning plant performance and distribution under changing climate regimes.

There are still major gaps in our understanding of rangeland degradation. Assessing the interactions between climate and grazing effects could help to explain what unifies and separates rangelands, and may therefore promote a more sustainable management of livestock. We studied 15 local land-use transects along a 600 km long climatic gradient in Central Asia to test the hypothesis that grazing effects differ between relatively moist equilibrium (EQ) and dry non-equilibrium (NEQ) rangeland systems. We analysed plant community composition, species diversity and indicator species for different grazing intensities. We found pronounced differences in community composition along our climate gradient, revealed climate-related grazing effects on richness, responses of Simpson's diversity, and also found different grazing indicator species along the larger transect. We conclude that in NEQ rangelands, grazing effects are limited to sacrifice zones and environmental filtering dominates vegetation composition. With increasing precipitation, resource availability gains in importance leading to more complex communities dominated by grazing-tolerant species under EQ dynamics. Hints for xerophytization in the transition zone between EQ and NEQ highlight the vulnerability of rangelands that temporally shift from one state to the other. This calls for extra care in the management of livestock numbers in these transition areas.

Based on fossil organism remains including plant macrofossils, charcoal, pollen, and invertebrates preserved in syngenetic deposits of the Batagay permafrost sequence in the Siberian Yana Highlands, we reconstructed the environmental history during marine isotope stages (MIS) 6 to 2. Two fossil assemblages, exceptionally rich in plant remains, allowed for a detailed description of the palaeo-vegetation during two climate extremes of the Late Pleistocene, the onset of the last glacial maximum (LGM) and the last interglacial. In addition, altogether 41 assemblages were used to outline the vegetation history since the penultimate cold stage of MIS 6. Accordingly, meadow steppes analogue to modern communities of the phytosociological order Festucetalia lenensis formed the primary vegetation during the Saalian and Weichselian cold stages. Cold-resistant tundra-steppe communities (Carici rupestris-Kobresietea bellardii) as they occur above the treeline today were, in contrast to more northern locations, mostly lacking. During the last interglacial, open coniferous woodland similar to modern larch taiga was the primary vegetation at the site. Abundant charcoal indicates wildfire events during the last interglacial. Zoogenic disturbances of the local vegetation were indicated by the presence of ruderal plants, especially by abundant Urtica dioica, suggesting that the area was an interglacial refugium for large herbivores. Meadow steppes, which formed the primary vegetation during cold stages and provided potentially suitable pastures for herbivores, were a significant constituent of the plant cover in the Yana Highlands also under the full warm stage conditions of the last interglacial. Consequently, meadow steppes occurred in the Yana Highlands during the entire investigated timespan from MIS 6 to MIS 2 documenting a remarkable environmental stability. Thus, the proportion of meadow steppe vegetation merely shifted in response to the respectively prevailing climatic conditions. Their persistence indicates low precipitation and a relatively warm growing season throughout and beyond the late Pleistocene. The studied fossil record also proves that modern steppe occurrences in the Yana Highlands did not establish as late as in the Holocene but instead are relicts of a formerly continuous steppe belt extending from Central Siberia to Northeast Yakutia during the Pleistocene. The persistence of plants and invertebrates characteristic of meadow steppe vegetation in interior Yakutia throughout the late Quaternary indicates climatic continuity and documents the suitability of this region as a refugium also for other organisms of the Pleistocene mammoth steppe including the iconic large herbivores. (C)2018 Elsevier Ltd. All rights reserved.

Restoration success of species-rich grasslands is often limited by low seed dispersal. To reintroduce target species of local provenances, bulk topsoil transfer is performed by excavating soil and vegetation from donor sites and spreading these on receptor sites. Our first objective was to determine the most advantageous season for transferring topsoil in Mediterranean grasslands. The second objective was to assess which treatmentcombination of season (spring or autumn) and transfer ratio (1:1 or 1:3)-performed best in restoring a Mediterranean grassland through bulk topsoil transfer after two large restoration projects. Just-in-time soil transfer (i.e. with no stockpiling) was implemented: (1) on a former 3ha orchard where topsoil was spread after removing trees and leveling soil; and (2) after a 5ha oil pipeline leak where polluted soil was removed prior to treatment (soil horizons were reconstituted). A seed bank study showed that the summer seed bank contained higher seed densities, species richness, and similarity to the reference site than the spring seed bank. Spring transfers gave better results than autumn transfers in terms of species richness and composition similarity with the reference site, while transfer ratios gave similar results. Long-term success was not driven by season or transfer ratio but by the underlying seed bank at receptor sites: the former orchard's weed-containing seed bank hampered topsoil transfer success. This study also suggests that restoration success cannot be deduced from seed bank studies alone, as species establishment is highly dependent on differences in growing conditions (including competition at receptor sites).

Phenological responses to changing temperatures are known as “fingerprints of climate change,” yet these reactions are highly species specific. To assess whether different plant characteristics are related to these species‐specific responses in flowering phenology, we observed the first flowering day (FFD) of ten herbaceous species along two elevational gradients, representing temperature gradients. On the same populations, we measured traits being associated with (1) plant performance (specific leaf area), (2) leaf biochemistry (leaf C, N, P, K, and Mg content), and (3) water‐use efficiency (stomatal pore area index and stable carbon isotopes concentration). We found that as elevation increased, FFD was delayed for all species with a highly species‐specific rate. Populations at higher elevations needed less temperature accumulation to start flowering than populations of the same species at lower elevations. Surprisingly, traits explained a higher proportion of variance in the phenological data than elevation. Earlier flowering was associated with higher water‐use efficiency, higher leaf C, and lower leaf P content. In addition to that, the intensity of shifts in FFD was related to leaf N and K. These results propose that traits have a high potential in explaining phenological variations, which even surpassed the effect of temperature changes in our study. Therefore, they have a high potential to be included in future analyses studying the effects of climate change and will help to improve predictions of vegetation changes.

Stomata enable plants to balance uptake of CO 2 with water loss via transpiration. Previous studies have shown that stomatal size and density trade-off with one another and are related to a suite of environmental factors (light, atmospheric CO 2 concentrations, moisture and temperature) which influence gas exchange. We examined the hypothesis that the habitat preferences of species are reflected in stomatal traits. Thus, we examined the relation of stomatal size and density of 36 herbaceous species along elevational gradients in relation to Ellenberg indicator values for light and moisture. The light value is a proxy for the transpiration demand due to exposure to radiation and air mass exchange, the moisture value reflects water supply. Additionally we measured carbon isotope discrimination (Δ 13 C), a proxy for intrinsic water use efficiency and internal CO 2 concentration. Stomatal size changed in parallel on both sides of a leaf, whereas changes in density differed between sides depending on species identity. There was an increase in absolute variation of sizes and densities with increasing mean size and density respectively, but not in relative variation. Species with few but large stomata tended to adjust stomatal size across environmental gradients, whereas species with small but many stomata mainly adjusted stomatal density. A higher Ellenberg indicator value for light and a lower value for moisture was associated with equal distribution of stomata between leaf sides. The carbon isotope discrimination data indicated that amphistomatic species, which illustrated a preference for high radiation and high air mass exchange as well as for dry habitats, had higher water use efficiency. We conclude that stomatal traits such as size, density or the distribution between the two sides of the leaf are indicators of how species optimize carbon uptake and balance water loss and radiation gain.

In dry steppes, strong climatic constraints, especially highly variable precipitation, and grazing are the most important factors controlling plant life. Growth is strongly limited by water availability, while grazing may affect species presence and performance. However, there is a lack of studies on population genetics of dryland plants in general, and of those addressing grazing effects in particular. To determine the landscape-scale genetic structure of dryland species, and if grazing has an impact on that, we chose the Eurasian steppe grass Stipa glareosa for a population genetic study employing nine polymorphic Simple Sequence Repeat (SSR) markers. We assessed genetic fingerprints of 200 individuals from six populations in Mongolia, which were sampled along a large-scale precipitation and altitudinal gradient. Nested within this gradient, sub-populations were sampled along short local transects representing different grazing intensities. Overall, S. glareosa populations showed rather low levels of genetic diversity at a mean Bruvo distance among individuals within a given population of 0.494 (mean expected heterozygosity He = 0.053). Linear mixed model analysis implied that genetic diversity was affected by both climatic constraints and local grazing conditions. We found a moderate isolation-by-distance pattern across all populations; grazing additionally influenced the genetic structure at local scale. Analysis of Molecular Variance revealed a modest genetic differentiation between populations (9 % of variation) and among sub-populations representing different grazing levels (11 %). Moreover, we detected indicator alleles that were exclusive for populations along the precipitation gradient; other alleles were associated with certain grazing levels across all populations. Thus, our data suggest that climatic constraints affect the genetic structure of S. glareosa populations, while at local scales differences in grazing disturbance may also matter.

Aim: Numerous studies have reported changes in first flowering day (FFD-changes) in response to changes in climate. However, regarding the direction (advances versus delays) and the intensity (number of days/decade) of FFD-changes, species show differences even when observed in the same location. Here, we examine the extent to which plant traits can explain observed differences in the response of flowering phenology in trees, shrubs, herbs and grasses. Location: Eighteen sites distributed over the Northern Hemisphere. Methods: We compiled data from the literature on FFD-changes over recent decades for 562 species (648 observations). We related FFD-changes to predictor variables associated with (a) changes in climate, (b) local site conditions and (c) traits. Results: Of all FDD-changes, 80.4% were FFD-advances, 69.9% not exceeding 5 days/decade, and 10.5% exceeding 5 days/decade, whereas 19.6% reported delays. The intensity of FFD-advances could be explained by several predictor variables from all three groups (a–c). The importance of these variables differed between the growth forms. Overall, decreasing precipitation was more important than increasing temperature in explaining FFD-advances. FFD-advances were strongest in polar tundra and in dry and warm habitats. Traits related to competition and growth rate, like plant height, specific leaf area and leaf dry matter content, had substantial explanatory power in the models. Traits had the highest overall importance in trees and grasses. In herbs they were of equal importance with changes in climate. In shrubs, variables related to site conditions best explained the intensity of FFD-advances. Main conclusions: Plant traits are important to understand species-specific and growth form-specific differences in phenological responses to climatic changes. Hence, in future observations and predictions of plant phenology, traits should be taken into consideration, especially those related to competition and growth rate, as they improve our understanding of adaptations leading to phenological changes.

International audience; Plant functional traits directly affect ecosystem functions. At the species level, trait combinations depend on trade-offs representing different ecological strategies, but at the community level trait combinations are expected to be decoupled from these trade-offs because different strategies can facilitate co-existence within communities. A key question is to what extent community-level trait composition is globally filtered and how well it is related to global versus local environmental drivers. Here, we perform a global, plot-level analysis of trait-environment relationships, using a database with more than 1.1 million vegetation plots and 26,632 plant species with trait information. Although we found a strong filtering of 17 functional traits, similar climate and soil conditions support communities differing greatly in mean trait values. The two main community trait axes that capture half of the global trait variation (plant stature and resource acquisitiveness) reflect the trade-offs at the species level but are weakly associated with climate and soil conditions at the global scale. Similarly, within-plot trait variation does not vary systematically with macro-environment. Our results indicate that, at fine spatial grain, macro-environmental drivers are much less important for functional trait composition than has been assumed from floristic analyses restricted to co-occurrence in large grid cells. Instead, trait combinations seem to be predominantly filtered by local-scale factors such as disturbance, fine-scale soil conditions, niche partitioning and biotic interactions.

Although specific leaf area (SLA) has been proposed to reflect plant responses to climatic changes, the link between SLA and temperature has never been systematically evaluated. Using in situ measured SLA values for 223 species occurring in 29 calcareous grasslands along a temperature gradient in the Bavarian Alps, we explored the SLA–temperature relationship at population (intraspecific), species (interspecific) and community level and investigated the relative impact of other environmental factors on SLA variation along the temperature gradient at the community level. Only 14 % of the studied species showed significant changes in their SLA values along the temperature gradient, despite high intraspecific variability of the SLA values. At the species level, we revealed a very weak positive SLA–temperature relationship (r 2 = 0.04, p

Summary Plant communities and their ecosystem functions are expected to be more resilient to future habitat fragmentation and deterioration if the species comprising the communities have a wide range of dispersal and persistence strategies. However, the extent to which the diversity of dispersal and persistence traits in plant communities is determined by the current and historical characteristics of sites and their surrounding landscape has yet to be explored. Using quantitative information on long-distance seed dispersal potential by wind and animals (dispersal in space) and on species' persistence/longevity (dispersal in time), we (i) compared levels of dispersal and persistence trait diversity (functional richness, FRic, and functional divergence, FDiv) in seminatural grassland plant communities with those expected by chance, and (ii) quantified the extent to which trait diversity was explained by current and historical landscape structure and local management history – taking into account spatial and phylogenetic autocorrel. Null model analysis revealed that more grassland communities than expected had a level of trait diversity that was lower or higher than predicted, given the level of species richness. Both the range (FRic) and divergence (FDiv) of dispersal and persistence trait values increased with grassland age. FDiv was mainly explained by the interaction between current grazing intensity and the amount of grassland habitat in the surrounding landscape in 1938. Synthesis. The study suggests that the variability of dispersal and persistence traits in grassland plant communities is driven by deterministic assembly processes, with both history and current management (and their interactions), playing a major role as determinants of trait diversity. While a long continuity of grazing management is likely to have promoted the diversity of dispersal and persistence traits in present-day grasslands, communities in sites that are well grazed at the present day, and were also surrounded by large amounts of grassland in the past, showed the highest diversity of dispersal and persistence strategies. Our results indicate that the historical context of a site within a landscape will influence the extent to which current grazing management is able to maintain a diversity of dispersal and persistence strategies and buffer communities (and their associated functions) against continuing habitat fragmentation. (Less)

In European dry grasslands land-use changes affect plant species performance and frequency. Potential driving forces are eutrophication and habitat fragmentation. The importance of these factors is presumably scale dependent. We used a functional trait approach to detect processes that influence species frequency and endangerment on different spatial scales. We tested for associations between functional traits and (1) frequency and (2) degree of endangerment on local, regional and national scales. We focussed on five selected traits that describe the life-history of plant species and that are related to competition, dispersal ability and habitat specificity. Trait data on plant height, SLA, plant coverage, peak of flowering and diaspore mass were measured for 28 perennials from common to rare and endangered to non-endangered on 59 dry grassland sites in north-eastern Germany. Multiple regression models revealed that species frequency is positively and species endangerment negatively related to plant height, plant coverage and SLA on more than one spatial scale. On the local scale, diaspore mass has a negative effect on species frequency. More frequent and less endangered species show a later peak of flowering on nationwide and regional scales. We concluded that competition traits are more important on larger scales, whereas dispersal traits are more important for species frequency on the smaller scale. On national and regional scales, eutrophication and habitat loss may be the main drivers of species threat, whereas on the local scale fragmentation plays a crucial role for the performance of dry grassland species.

Why species are found where they are is a central question in biogeography. The most widely used tool for understanding the controls on distribution is species distribution modelling. Species distribution modelling is now a well-established method in both the theoretical and applied ecological literature. In this special issue we examine the current state of the art in species distribution modelling and explore avenues for including more biological processes in such models. In particular we focus on physiological, demographic, dispersal, competitive and ecological-modulation processes. This overview highlights opportunities for new species distribution model concepts and developments, as well as a statistical agenda for implementing such models.

Within the field of species distribution modelling an apparent dichotomy exists between process-based and correlative approaches, where the processes are explicit in the former and implicit in the latter. However, these intuitive distinctions can become blurred when comparing species distribution modelling approaches in more detail. In this review article, we contrast the extremes of the correlative–process spectrum of species distribution models with respect to core assumptions, model building and selection strategies, validation, uncertainties, common errors and the questions they are most suited to answer. The extremes of such approaches differ clearly in many aspects, such as model building approaches, parameter estimation strategies and transferability. However, they also share strengths and weaknesses. We show that claims of one approach being intrinsically superior to the other are misguided and that they ignore the process–correlation continuum as well as the domains of questions that each approach is addressing. Nonetheless, the application of process-based approaches to species distribution modelling lags far behind more correlative (process-implicit) methods and more research is required to explore their potential benefits. Critical issues for the employment of species distribution modelling approaches are given, together with a guideline for appropriate usage. We close with challenges for future development of process-explicit species distribution models and how they may complement current approaches to study species distributions.

Aim Biotic interactions – within guilds or across trophic levels – have widely been ignored in species distribution models (SDMs). This synthesis outlines the development of ‘species interaction distribution models’ (SIDMs), which aim to incorporate multispecies interactions at large spatial extents using interaction matrices. Location Local to global. Methods We review recent approaches for extending classical SDMs to incorporate biotic interactions, and identify some methodological and conceptual limitations. To illustrate possible directions for conceptual advancement we explore three principal ways of modelling multispecies interactions using interaction matrices: simple qualitative linkages between species, quantitative interaction coefficients reflecting interaction strengths, and interactions mediated by interaction currencies. We explain methodological advancements for static interaction data and multispecies time series, and outline methods to reduce complexity when modelling multispecies interactions. Results Classical SDMs ignore biotic interactions and recent SDM extensions only include the unidirectional influence of one or a few species. However, novel methods using error matrices in multivariate regression models allow interactions between multiple species to be modelled explicitly with spatial co-occurrence data. If time series are available, multivariate versions of population dynamic models can be applied that account for the effects and relative importance of species interactions and environmental drivers. These methods need to be extended by incorporating the non-stationarity in interaction coefficients across space and time, and are challenged by the limited empirical knowledge on spatio-temporal variation in the existence and strength of species interactions. Model complexity may be reduced by: (1) using prior ecological knowledge to set a subset of interaction coefficients to zero, (2) modelling guilds and functional groups rather than individual species, and (3) modelling interaction currencies and species’ effect and response traits. Main conclusions There is great potential for developing novel approaches that incorporate multispecies interactions into the projection of species distributions and community structure at large spatial extents. Progress can be made by: (1) developing statistical models with interaction matrices for multispecies co-occurrence datasets across large-scale environmental gradients, (2) testing the potential and limitations of methods for complexity reduction, and (3) sampling and monitoring comprehensive spatio-temporal data on biotic interactions in multispecies communities.

Although urban areas only occupy c. 2.8% of the earth's land surface, urbanization threatens biodiversity as areas of high human population density often coincide with high biodiversity. Therefore, nature conservation should concentrate on both remote areas and densely populated regions. Protecting rare plant species in rural and urban areas can contribute to the protection of biodiversity. We therefore need to understand why species are rare. Studies on causes of rarity often concentrate on either plant traits or extrinsic threats (such as habitat fragmentation or nitrogen enrichment). However, there are only a few studies that combine traits and extrinsic threats, although such analyses might clarify causes of rarity. We assessed how the affinity of vascular plant species to urban land use ('urbanity') interacts with plant traits in determining species frequency.Germany, resolution c. 12 km x 11 km.Species with a low frequency may be rare because they occur in rare habitats or because of other reasons, although their habitat is frequent. Therefore, we calculated the frequency of species corrected for habitat frequency, i.e. relative species frequency. We explained relative species frequency by the interactions of species traits and species affinity to urban land use using generalized linear models. Simultaneous autoregressive error models controlled for phylogenetic relationships of species.Relative species frequency depends on species affinity to urban land use, independent of the different interactions between traits and urbanity used as predictors. The higher the urbanity the higher is species frequency. Urbanity interacts with species preferences towards temperature and soil acidity. Moreover, dispersal, nitrogen preferences and origin explain relative species frequency, amongst others.Many rare species, especially those preferring cool or acidic habitats might already have disappeared from urban areas. Analyses that combine species traits and environmental effects can explain the causes of rarity and help to derive better conservation strategies.

International audience; The ongoing decline of many plant species in Northwest Europe indicates that traditional conservation measures to improve the habitat quality, although useful, are not enough to halt diversity losses. Using recent databases, we show for the first time that differences between species in adaptations to various dispersal vectors, in combination with changes in the availability of these vectors, contribute significantly to explaining losses in plant diversity in Northwest Europe in the 20th century. Species with water- or fur-assisted dispersal are over-represented among declining species, while others (windor bird-assisted dispersal) are under-represented. Our analysis indicates that the 'colonization deficit' due to a degraded dispersal infrastructure is no less important in explaining plant diversity losses than the more commonly accepted effect of eutrophication and associated niche-based processes. Our findings call for measures that aim to restore the dispersal infrastructure across entre regions and that go beyond current conservation practices.

Within the past few years plant functional trait analyses have been widely applied to learn more about the processes and patterns of ecosystem development in response to environmental changes. These approaches are based on the assumption that plants with similar ecologically relevant trait attributes respond to environmental changes in comparable ways. Several methods have been described on how to analyse a priori defined trait sets with respect to environment. Irrespective of the statistical methods used to contrast ecosystem responses and environmental conditions, each functional trait approach depends strongly on the initial trait set. In nearly all recent studies on functional trait analysis a test, if a trait is responsible, is applied independently from the core analysis. In the current study we present a method that extracts those traits from a wider set of traits which are optimal for describing the ecosystem response to a given environmental gradient. This was done by the use of iterative three-table ordination techniques with each possible trait combination. We further concentrated on the effect of the inclusion of too many traits in such analyses. As examples the method was applied to three long term studies on abandoned arable fields. The approach was validated by comparing the results with literature-knowledge on arable field succession. Although the trait pre-selection was only based on a statistical procedure, our method was able to identify all relevant processes of ecosystem responses. All three sites show comparable ecosystem responses; the importance of the competitive ability of plants was highlighted. We further demonstrated that the use of too many traits results in an over-fitting of the trait-environment model. The presented method of iterative RLQ-analyses is adequate to identify responding traits to environmental changes: the discovered processes of successional development of abandoned arable fields are consistent with our knowledge from the literature.

1. An international group of scientists has built an open internet data base of life-history traits of the Northwest European flora (the LEDA-Traitbase) that can be used as a data source for fundamental research on plant biodiversity and coexistence, macro-ecological patterns and plant functional responses.2. The species-trait matrix comprises referenced information under the control of an editorial board, for ca. 3000 species of the Northwest European flora, combining existing information and additional measurements. The data base currently contains data on 26 plant traits that describe three key features of plant dynamics: persistence, regeneration and dispersal. The LEDA-Traitbase is freely available at www.leda-traitbase.org.3. We present the structure of the data base and an overview of the trait information available.4. Synthesis. The LEDA Traitbase is useful for large-scale analyses of functional responses of communities to environmental change, effects of community trait composition on ecosystem properties and patterns of rarity and invasiveness, as well as linkages between traits as expressions of fundamental trade-offs in plants.

Due to ubiquitous eutrophication and fragmentation, many plant species are actually threatened in Europe. Most ecosystems face an overall nutrient input leading to changes in species composition. Fragmentation is effectively influencing species survival. We investigate if two different measures of species performance of 91 calcareous grassland species–rate of decline and rarity—are related to comparable traits and hence processes. On the one hand we expected that species rate of decline is mainly determined by the processes of eutrophication and fragmentation. On the other hand we hypothesized that the importance of site characteristics may overwhelm the effect of eutrophication and fragmentation for species rarity. Hence, we compared persistence traits responding to eutrophication, dispersal traits being related to fragmentation and ecological site factors for decreasing and increasing species and for rare and common species. The results suggest that increasing species had better means of long-distance dispersal and were more competitive than decreasing species. In contrast, there were hardly any differences in traits between rare and common species, but site characteristics were related to species rarity. Rare species were in the main those with ecological preferences for warm, dry, light and nutrient poor conditions. This study may represent a basis for the assessment of plant species threat. Applying the deduced knowledge about the life history of decreasing versus increasing species to habitat-scale approaches it is possible to predict which species may become threatened in the future simply from the combination of their trait values.

Aim Species frequency data have been widely used in nature conservation to aid management decisions. To determine species frequencies, information on habitat occurrence is important: a species with a low frequency is not necessarily rare if it occupies all suitable habitats. Often, information on habitat distribution is available for small geographic areas only. We aim to predict grid-based habitat occurrence from grid-based plant species distribution data in a meso-scale analysis. Location The study was carried out over two spatial extents: Germany and Bavaria. Methods Two simple models were set up to examine the number of characteristic plant species needed per grid cell to predict the occurrence of four selected habitats (species data from FlorKart, http://www.floraweb.de). Both models were calibrated in Bavaria using available information on habitat distribution, validated for other federal states, and applied to Germany. First, a spatially explicit regression model (generalized linear model (GLM) with assumed binomial error distribution of response variable) was obtained. Second, a spatially independent optimization model was derived that estimated species numbers without using spatial information on habitat distribution. Finally, an additional uncalibrated model was derived that calculated the frequencies of 24 habitats. It was validated using NATURA2000 habitat maps.

It is generally assumed that morphological structures like awns, bristles, or hooks increase retention potential of plant diaspores to the coats of animals, but this basic assumption has rarely been tested. In this paper we introduce a lab-experiment in which detachment rates of diaspores from mechanically shaken animal coats are measured using a standardized protocol and compared with detachment rates measured under natural conditions. The lab-experiments were used to assess ‘retention potentials’ of diaspores for more than 100 plant species. ‘Retention potential’ is defined as the proportion of diaspores still attached to the animal coat after a certain time period. We assessed the effect of several diaspore traits on retention potential and found that a strong negative effect of diaspore mass overrides the expected positive effect of structures like awns, bristles or hooks. However, a positive effect of these structures was proven, when analysing the dataset separately for diaspores of different mass classes. Additionally, we found a negative effect of flat appendages. Coat type also affects retention potentials, with higher values in curly sheep wool compared to straight cattle hair. Species with diaspore mass 2 mg might also be dispersed over long distances, at least by sheep. The frequency distribution of retention potentials clearly shows that a binary classification of species dispersed in animal hair vs. species not dispersed is artificial since differences are gradual. The relative importance of diaspore mass for determining retention potential was higher in straight hair, whereas diaspore morphology had a relatively higher impact in curly wool. Hence, different sets of plant species are dispersed by different animals.

Dispersal is a process that determines many aspects in the life-history of plants. Up to now, however, it is difficult to quantify. Many studies rather assess it as a categorical trait, i.e. assuming a species is dispersed by a certain vector or not. Gradual differences in the dispersal potential between species are rarely considered. In this paper we focus on the key process of epizoochory: the attachment potential of species to animal coats. We present two simple models (GLMs) how to quantify and to predict attachment potentials to sheep wool and to cattle hair from easily measurable seed traits. To calibrate the models, we investigated the attachment potential of seeds of 130 plant species on mechanically shaken coats of sheep and cattle and measured traits describing the seeds. We found that seed mass and seed morphology were correlated with attachment potentials. For sheep wool, the combination of the logarithm of seed mass and a constant for the different seed morphology types explained 85% of the variation of attachment potentials. For cattle hair, 71% were explained. To validate the models, they were applied to 36 additional species. Predicted and experimentally measured attachment potentials were correlated with r=0.84 for sheep wool and r=0.61 for cattle wool. Thus, the attachment potential of seeds to sheep or cattle coat can be assessed for a large set of species using the described models if seed mass and seed morphology are known.

In a recent article (Dormann et al., 2012, Journal of Biogeography, 39, 2119–2131), we compared different approaches to species distribution modelling and depicted modelling approaches along an axis from purely ‘correlative’ to ‘forward process-based’ models. In their correspondence, Kriticos et al. (2013, Journal of Biogeography, doi:10.1111/j.1365-2699.2012.02791.x) challenge this view, claiming that our continuum representation neglects differences among models and does not consider the ability of fitted process-based models to combine the advantages of both process-based and correlative modelling approaches. Here we clarify that the continuum view resulted from recognition of the manifold differences between models. We also reinforce the point that the current trend towards combining different modelling approaches may lead not only to the desired combination of the advantages but also to the accumulation of the disadvantages of those approaches. This point has not been made sufficiently clear previously.

In Europe, the present landscape has been mainly influenced by human activities. Especially for dry semi-natural grasslands, agricultural intensification led to a considerable habitat loss and fagmentation. In the Crau, large parts of the former 60,000 ha semi-natural steppe vegetation (coussous) have been transformed into arable, industrial and military land during the 20th century. However, in the 1980s, most of the cereal and melon fields have been abandoned and sheep grazing has been reintroduced. The current study investigates the effects of historical melon and cereal cultivation on the steppe vegetation and the potential of characteristic coussous species to re-establish after abandonment. At four sites of central Crau, vegetation sampling has been carried out on the different vegetation types (coussous, melon and cereal fllows) using the frequency method. Soil samples were taken and analysed. At one site, seed bank studies were performed. As related to seed bank persistence, dormancy traits were examined for 26 characteristic steppe species. Decades after abandonment, the vegetation of coussous, abandoned cereal and abandoned melon fields are still very different from each other : Due to former fertiliser application, ex-cultivated cereal and melon fields still showed enhanced nutrient levels (N, P, K, Ca, pH). The seed bank of the coussous is dominated by transient species while abandoned field seed banks are dominated by persistent seeds of ruderal species. The germination studies confirmed the absence of dormancy mechanisms for seeds of most coussous species. The successful reestablishment of coussous species in abandoned fields seemed to be hampered by still modified abiotic conditions. More, most coussous species do not build up a persistent soil seed bank and can, therefore, not survive cultivation periods. Species dispersal between coussous and adjacent ex-cultivated fields appeared to be a limiting fctor, too. For following studies complete species lists of ex-cultivated fields and the coussous are provided, En Europe, les paysages actuels ont été en grande partie influencés par les activités humaines. Ainsi, pour les paysages de pelouses sèches, l'intensification des pratiques agricoles a entraîné une perte considérable et une fragmentation importante des habitats. Dans la plaine de Crau, de grandes zones de la steppe semi-naturelle (coussous) ont été transformées en surfaces cultivées, industrielles et militaires durant le XXe siècle. Cependant, dans les années 80, la plupart des cultures de céréales et de melons ont été abandonnées au profit d'un retour au pâturage ovin. Le présent travail a pour objectif de rechercher, d'une part, les effets des cultures de melons et de céréales anciennes sur la végétation steppique et, d'autre part, les potentialités de ré-établissement dans les fiches des espèces steppiques les plus caractéristiques. Dans quatre sites du centre de la Crau, la végétation a été échantillonnée dans différents types de parcelles (coussous, fiches melonnières et céréalières) par la méthode des quadrats. Des échantillons de sol ont également été prélevés et analysés. Pour un site, la banque de graines a été étudiée. En liaison avec la viabilité de la banque de graines dans le sol, les traits biologiques relatifs à la dormance ont été examinés pour 26 espèces caractéristiques de la steppe. Quelques décennies après l'abandon cultural, la végétation des coussous, fiches melonnières et céréalières est très différente. Suite à l'épandage d'engrais et à des labours profonds, les fiches melonnières et céréalières possèdent des teneurs supérieures en éléments minéraux (N, P205, K2 0, CaO). La banque de graines du coussous est dominée par des espèces à viabilité transitoire ( 5 ans) d'espèces rudérales. Les tests de germination confirment l'absence de dormance pour la plupart des espèces du coussous. Le ré-établissement des espèces du coussous dans les champs abandonnés semble ralenti par des changements dans les conditions abiotiques. La plupart des espèces du coussous ne possèdent pas de banque de graines, et donc, ces espèces ne peuvent pas survivre à des périodes de cultures. Les processus de dissémination des espèces entre les coussous et les champs anciennement cultivés apparaissent également comme un facteur limitant. Pour des études futures, des listes complètes d'espèces des fiches et des coussous sont fournies en annexes., Römermann Christine, Bernhardt Markus, Dutoit Thierry, Poschlod Peter, Rolando Christiane. Histoire culturale de la Crau : potentialités de ré-établissement des espèces caractéristiques du coussous après abandon. In: Ecologia mediterranea, tome 30 n°1, 2004. Biologie de la conservation et gestion des espaces naturels en Crau / Biological conservation and management of the natural areas in the Crau. pp. 47-70.

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.