There is growing recognition that classifying terrestrial plant species on the basis of their function (into 'functional types') rather than their higher taxonomic identity, is a promising way forward for tackling important ecological questions at the scale of ecosystems, landscapes or biomes. These questions include those on vegetation responses to and vegetation effects on, environmental changes (e.g. changes in climate, atmospheric chemistry, land use or other disturbances). There is also growing consensus about a shortlist of plant traits that should underlie such functional plant classifications, because they have strong predictive power of important ecosystem responses to environmental change and/or they themselves have strong impacts on ecosystem processes. The most favoured traits are those that are also relatively easy and inexpensive to measure for large numbers of plant species. Large international research efforts, promoted by the IGBP–GCTE Programme, are underway to screen predominant plant species in various ecosystems and biomes worldwide for such traits. This paper provides an international methodological protocol aimed at standardising this research effort, based on consensus among a broad group of scientists in this field. It features a practical handbook with step-by-step recipes, with relatively brief information about the ecological context, for 28 functional traits recognised as critical for tackling large-scale ecological questions.

The authors regret that elements of Appendix 1 were incorrect in the original publication. The correct version of Appendix 1 is given below. Appendix 1. Summary of plant traits Summary of plant traits included in the handbookThe range of values corresponds to those generally reported for field-grown plants. Ranges of values are based on the literature and the authors' datasets and do not always necessarily correspond to the widest ranges that exist in nature or are theoretically possible. Recommended sample size indicates the minimum and preferred number of individuals to be sampled, so as to obtain an appropriate indication of the values for the trait of interest; when only one value is given, it corresponds to the number of individuals ( = replicates); when two values are given, the first one corresponds to the number of individuals and the second one to the number of organs to be measured per individual. Note that one replicate can be compounded from several individuals (for smaller species), whereas one individual cannot be used for different replicates. The expected coefficient of variation (CV) range gives the 20th and the 80th percentile of the CV ( = s.d. scaled to the mean) as observed in several datasets obtained for a range of field plants for different biomes. Numbering of plant traits corresponds with the numbering of the chapters in the handbook Abstract. Plant functional traits are the features (morphological, physiological, phenological) that represent ecological strategies and determine how plants respond to environmental factors, affect other trophic levels and influence ecosystem properties. Variation in plant functional traits, and trait syndromes, has proven useful for tackling many important ecological questions at a range of scales, giving rise to a demand for standardised ways to measure ecologically meaningful plant traits. This line of research has been among the most fruitful avenues for understanding ecological and evolutionary patterns and processes. It also has the potential both to build a predictive set of local, regional and global relationships between plants and environment and to quantify a wide range of natural and human-driven processes, including changes in biodiversity, the impacts of species invasions, alterations in biogeochemical processes and vegetation-atmosphere interactions. The importance of these topics dictates the urgent need for more and better data, and increases the value of standardised protocols for quantifying trait variation of different species, in particular for traits with power to predict plant-and ecosystemlevel processes, and for traits that can be measured relatively easily. Updated and expanded from the widely used previous version, this handbook retains the focus on clearly presented, widely applicable, step-by-step recipes, with a minimum of text on theory, and not only includes updated methods for the traits previously covered, but also introduces many new protocols for further traits. This new handbook has a better balance between whole-plant ...

This paper discusses the advantages and disadvantages of the different methods that separate net ecosystem exchange (NEE) into its major components, gross ecosystem carbon uptake (GEP) and ecosystem respiration (R-eco). In particular, we analyse the effect of the extrapolation of night-time values of ecosystem respiration into the daytime; this is usually done with a temperature response function that is derived from long-term data sets. For this analysis, we used 16 one-year-long data sets of carbon dioxide exchange measurements from European and US-American eddy covariance networks. These sites span from the boreal to Mediterranean climates, and include deciduous and evergreen forest, scrubland and crop ecosystems. long-term (annual) data sets, does not reflect the short-term temperature sensitivity that is effective when extrapolating from night- to daytime. Specifically, in summer active ecosystems the long-term temperature sensitivity exceeds the short-term sensitivity. Thus, in those ecosystems, the application of a long-term temperature sensitivity to the extrapolation of respiration from night to day leads to a systematic overestimation of ecosystem respiration from half-hourly to annual time-scales, which can reach > 25% for an annual budget and which consequently affects estimates of GEP. Conversely, in summer passive (Mediterranean) ecosystems, the long-term temperature sensitivity is lower than the short-term temperature sensitivity resulting in underestimation of annual sums of respiration. temperature sensitivity of R-eco from eddy covariance data that applies this to the extrapolation from night- to daytime, and that further performs a filling of data gaps that exploits both, the covariance between fluxes and meteorological drivers and the temporal structure of the fluxes. While this algorithm should give less biased estimates of GEP and R-eco, we discuss the remaining biases and recommend that eddy covariance measurements are still backed by ancillary flux measurements that can reduce the uncertainties inherent in the eddy covariance data. [References: 53

Future climate warming is expected to enhance plant growth in temperate ecosystems and to increase carbon sequestration. But although severe regional heatwaves may become more frequent in a changing climate, their impact on terrestrial carbon cycling is unclear. Here we report measurements of ecosystem carbon dioxide fluxes, remotely sensed radiation absorbed by plants, and country-level crop yields taken during the European heatwave in 2003. We use a terrestrial biosphere simulation model to assess continental-scale changes in primary productivity during 2003, and their consequences for the net carbon balance. We estimate a 30 per cent reduction in gross primary productivity over Europe, which resulted in a strong anomalous net source of carbon dioxide (0.5 Pg C yr(-1)) to the atmosphere and reversed the effect of four years of net ecosystem carbon sequestration. Our results suggest that productivity reduction in eastern and western Europe can be explained by rainfall deficit and extreme summer heat, respectively. We also find that ecosystem respiration decreased together with gross primary productivity, rather than accelerating with the temperature rise. Model results, corroborated by historical records of crop yields, suggest that such a reduction in Europe's primary productivity is unprecedented during the last century. An increase in future drought events could turn temperate ecosystems into carbon sources, contributing to positive carbon-climate feedbacks already anticipated in the tropics and at high latitudes.

Concern is growing about the consequences of biodiversity loss for ecosystem functioning, for the provision of ecosystem services, and for human well being. Experimental evidence for a relationship between biodiversity and ecosystem process rates is compelling, but the issue remains contentious. Here, we present the first rigorous quantitative assessment of this relationship through meta-analysis of experimental work spanning 50 years to June 2004. We analysed 446 measures of biodiversity effects (252 in grasslands), 319 of which involved primary producer manipulations or measurements. Our analyses show that: biodiversity effects are weaker if biodiversity manipulations are less well controlled; effects of biodiversity change on processes are weaker at the ecosystem compared with the community level and are negative at the population level; productivity-related effects decline with increasing number of trophic links between those elements manipulated and those measured; biodiversity effects on stability measures (ÔinsuranceÕ effects) are not stronger than biodiversity effects on performance measures. For those ecosystem services which could be assessed here, there is clear evidence that biodiversity has positive effects on most. Whilst such patterns should be further confirmed, a precautionary approach to biodiversity management would seem prudent in the meantime.

The respiratory activities of plant roots, of their mycorrhizal fungi and of the free-living microbial heterotrophs (decomposers) in soils are significant components of the global carbon balance, but their relative contributions remain uncertain. To separate mycorrhizal root respiration from heterotrophic respiration in aboreal pine forest, we conducted a large-scale tree-girdling experiment, comprising 9 plots each containing about 120 trees. Tree-girdling involves stripping the stem bark to the depth of the current xylem at breast height terminating the supply of current photosynthates to roots and their mycorrhizal fungi without physically disturbing the delicate root-microbe-soil system. Here we report that girdling reduced soil respiration within 1-2 months by about 54% relative to respiration on ungirdled control plots, and that decreases of up to 37% were detected within 5 days. These values clearly show that the flux of current assimilates to roots is a key driver of soil respiration; they are conservative estimates of root respiration, however, because girdling increased the use of starch reserves in the roots. Our results indicate that models of soil respiration should incorporate measures of photosynthesis and of seasonal patterns of photosynthate allocation to roots.

The terrestrial biosphere is a key component of the global carbon cycle and its carbon balance is strongly influenced by climate. Continuing environmental changes are thought to increase global terrestrial carbon uptake. But evidence is mounting that climate extremes such as droughts or storms can lead to a decrease in regional ecosystem carbon stocks and therefore have the potential to negate an expected increase in terrestrial carbon uptake. Here we explore the mechanisms and impacts of climate extremes on the terrestrial carbon cycle, and propose a pathway to improve our understanding of present and future impacts of climate extremes on the terrestrial carbon budget.

A; et al., (2010). Bottom-up effects of plant diversity on multitrophic interactions in a biodiversity experiment. Nature, (468) Bottom-up effects of plant diversity on multitrophic interactions in a biodiversity experiment Abstract Biodiversity is rapidly declining1, and this may negatively affect ecosystem processes, including economically important ecosystem services. Previous studies have shown that biodiversity has positive effects on organisms and processes4 across trophic levels. However, only a few studies have so far incorporated an explicit food-web perspective. In an eight-year biodiversity experiment, we studied an unprecedented range of above-and below-ground organisms and multitrophic interactions. A multitrophic data set originating from a single long-term experiment allows mechanistic insights that would not be gained from meta-analysis of different experiments. Here we show that plant diversity effects dampen with increasing trophic level and degree of omnivory. This was true both for abundance and species richness of organisms. Furthermore, we present comprehensive above-ground/below-ground biodiversity food webs. Both above ground and below ground, herbivores responded more strongly to changes in plant diversity than did carnivores or omnivores. Density and richness of carnivorous taxa was independent of vegetation structure. Below-ground responses to plant diversity were consistently weaker than above-ground responses. Responses to increasing plant diversity were generally positive, but were negative for biological invasion, pathogen infestation and hyperparasitism. Our results suggest that plant diversity has strong bottom-up effects on multitrophic interaction networks, with particularly strong effects on lower trophic levels. Effects on higher trophic levels are indirectly mediated through bottom-up trophic cascades. However, only few studies so far incorporated an explicit food-web perspective 6 . In an 8-50 year biodiversity experiment, we studied an unprecedented range of above-and 51 belowground organisms and biotic interactions. A multitrophic dataset originating from 52 a single long-term experiment allows mechanistic insights that otherwise would not be 53 possible using meta-analysis of different experiments. We show that plant diversity 54 effects dampen with increasing trophic level and degree of omnivory. This finding was 55 consistent both for abundance and species richness of organisms. Further, we present 56 comprehensive above-belowground biodiversity food webs. Both above-and 57 belowground, herbivores responded more strongly to changes in plant diversity than did 58 carnivores or omnivores. Density and richness of carnivorous taxa was independent of 59 vegetation structure. Belowground responses to plant diversity were consistently weaker 60 than aboveground responses. Responses to increasing plant diversity were generally 61 positive, but negative for biological invasion, pathogen infestation and hyperparasitism. 62Our results suggest that plant diversity exhibits stron...