Plant traits-the morphological, anatomical, physiological, biochemical and phenological characteristics of plants-determine how plants respond to environmental factors, affect other trophic levels, and influence ecosystem properties and their benefits and detriments to people. Plant trait data thus represent the basis for a vast area of research spanning from evolutionary biology, community and functional ecology, to biodiversity conservation, ecosystem and landscape management, restoration, biogeography and earth system modelling. Since its foundation in 2007, the TRY database of plant traits has grown continuously. It now provides unprecedented data coverage under an open access data policy and is the main plant trait database used by the research community worldwide. Increasingly, the TRY database also supports new frontiers of trait-based plant research, including the identification of data gaps and the subsequent mobilization or measurement of new data. To support this development, in this article we evaluate the extent of the trait data compiled in TRY and analyse emerging patterns of data coverage and representativeness. Best species coverage is achieved for categorical traits-almost complete coverage for 'plant growth form'. However, most traits relevant for ecology and vegetation modelling are characterized by continuous intraspecific variation and trait-environmental relationships. These traits have to be measured on individual plants in their respective environment. Despite unprecedented data coverage, we observe a humbling lack of completeness and representativeness of these continuous traits in many aspects.We, therefore, conclude that reducing data gaps and biases in the TRY database remains a key challenge and requires a coordinated approach to data mobilization and trait measurements. This can only be achieved in collaboration with other initiatives. Geosphere-Biosphere Program (IGBP) and DIVERSITAS, the TRY database (TRY-not an acronym, rather a statement of sentiment; https ://www.try-db.org; Kattge et al., 2011) was proposed with the explicit assignment to improve the availability and accessibility of plant trait data for ecology and earth system sciences. The Max Planck Institute for Biogeochemistry (MPI-BGC) offered to host the database and the different groups joined forces for this community-driven program. Two factors were key to the success of TRY: the support and trust of leaders in the field of functional plant ecology submitting large databases and the long-term funding by the Max Planck Society, the MPI-BGC and the German Centre for Integrative Biodiversity Research (iDiv) Halle-Jena-Leipzig, which has enabled the continuous development of the TRY database.
For the past century, woody plants have increased in grasslands and savannas worldwide. Woody encroachment may significantly alter ecosystem functioning including fire regimes, herbivore carrying capacity, biodiversity and carbon storage capacity. Traditionally, increases in woody cover and density have been ascribed to changes in the disturbance regime (fire and herbivores) or rainfall. Increased atmospheric CO 2 concentrations may also contribute, by increasing growth rates of trees relative to grasses. This hypothesis is still heavily debated because usually potential CO 2 effects are confounded by changes in land use (disturbance regime). Here we analyse changes in woody density in fire experiments at three sites in South African savannas where the disturbance regime (fire and herbivores) was kept constant for 30 and 50 years. If global drivers had significant effects on woody plants, we would expect significant increases in tree densities and biomass over time under the constant disturbance regime. Woody density remained constant in a semiarid savanna but tripled in a mesic savanna between the 1970s and 1990s. At the third site, a semiarid savanna near the southern limits of the biome, tree density doubled from the mid 1990s to 2010. Interpretation of the causes is confounded by population recovery after clearing, but aerial photograph analysis on adjacent non-cleared areas showed an accompanying 48% increase in woody cover. Increased CO 2 concentrations are consistent with increased woody density while other global drivers (rainfall) remained constant over the duration of the experiments. The absence of a response in one semiarid savanna could be explained by a smaller carbon sink capacity of the dominant species, which would therefore benefit less from increased CO 2 . Understanding how savannas and grasslands respond to increased CO 2 and identifying the causes of woody encroachment are essential for the successful management of these systems.
Changes in the phenology of vegetation activity may accelerate or dampen rates of climate change by altering energy exchanges between the land surface and the atmosphere 1,2 and can threaten species with synchronized life cycles 3-5 . Current knowledge of long-term changes in vegetation activity is regional 6-8 , or restricted to highly integrated measures of change such as net primary productivity 9-13 , which mask details that are relevant for Earth system dynamics. Such details can be revealed by measuring changes in the phenology of vegetation activity. Here we undertake a comprehensive global assessment of changes in vegetation phenology. We show that the phenology of vegetation activity changed severely (by more than 2 standard deviations in one or more dimensions of phenological change) on 54% of the global land surface between 1981 and 2012. Our analysis confirms previously detected changes in the boreal and northern temperate regions 6-8 . The adverse consequences of these northern phenological shifts for land-surface-climate feedbacks 1 , ecosystems 4 and species 3 are well known. Our study reveals equally severe phenological changes in the southern hemisphere, where consequences for the energy budget and the likelihood of phenological mismatches are unknown. Our analysis provides a sensitive and direct measurement of ecosystem functioning, making it useful both for monitoring change and for testing the reliability of early warning signals of change 14 .Recent climate change has shifted species distributions 15,16 and leaf phenology 17,18 around the world, leading to mismatches in previously synchronized phenological cycles 3,4 . Such mismatches greatly increase the risk of extinction for affected species, and ongoing climatic and phenological change is expected to further increase this risk 5 . Despite documenting and predicting effects of climate change on many organisms, these previous studies do not provide an easy way of inferring how widespread such changes are or where they are most severe. In addition to being a symptom of climate change, vegetation change also feeds back to the climate system by forcing rates of energy exchange between the land surface and the atmosphere. Changes in the vigour and timing of vegetation activity can therefore accelerate or slow down rates of climate change 1 . Yet, the extent to which changes in vegetation phenology will impact the climate system by modifying albedo, transpiration, partitioning between latent and sensible heat in the atmosphere, and cloud formation, has been identified as a major source of uncertainty in climate change projections 2,19 .
Biomes are important constructs for organizing understanding of how the worlds' major terrestrial ecosystems differ from one another and for monitoring change in these ecosystems. Yet existing biome classification schemes have been criticized for being overly subjective and for explicitly or implicitly invoking climate. We propose a new biome map and classification scheme that uses information on (i) an index of vegetation productivity, (ii) whether the minimum of vegetation activity is in the driest or coldest part of the year, and (iii) vegetation height. Although biomes produced on the basis of this classification show a strong spatial coherence, they show little congruence with existing biome classification schemes. Our biome map provides an alternative classification scheme for comparing the biogeochemical rates of terrestrial ecosystems. We use this new biome classification scheme to analyse the patterns of biome change observed over recent decades. Overall, 13% to 14% of analysed pixels shifted in biome state over the 30-year study period. A wide range of biome transitions were observed. For example, biomes with tall vegetation and minimum vegetation activity in the cold season shifted to higher productivity biome states. Biomes with short vegetation and low seasonality shifted to seasonally moisture-limited biome states. Our findings and method provide a new source of data for rigorously monitoring global vegetation change, analysing drivers of vegetation change and for benchmarking models of terrestrial ecosystem function.
Summary 1.The effects of moisture conditions, seed morphology, vegetation structure and hydrodynamic variables on seed retention were examined in a system where the dominant dispersal agent is water. Experiments were conducted in a tidal salt marsh and in a flume facility where hydrodynamic variables could be controlled. 2. Moisture condition of seeds greatly influenced which factors were most important in determining seed retention. Seed type (buoyancy) was the most important factor when seeds were dry with seeds possessing very low floating capacity ( Plantago maritima ) being retained in greater numbers than seeds with intermediate floating capacities ( Suaeda maritima and Elytrigia atherica ). 3. In contrast, hydrodynamic variables dominated retention processes when seeds were waterlogged. The application of waves in addition to flow velocity dislodged more seeds than flow velocity alone. 4. Vegetation structure influenced retention in both dry and wet conditions but less so than other factors. Denser, less rigid vegetation types retained greater numbers of seeds than more open, more rigid vegetation types. 5. Results suggest that buoyancy traits appear to determine whether seeds move in the drier summer and autumn months after initial detachment from parent plants but the intensity of wave action will determine whether waterlogged seeds stay in a microsite during the wetter months of late autumn to early spring.
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