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 ...

Vascular plants are the main entry point for energy and matter into the Earth's terrestrial ecosystems. Their Darwinian struggle for growth, survival and reproduction in very different arenas has resulted in an extremely wide variety of form and function, both across and within habitats. Yet it has long been thought 1-8 that there is a pattern to be found in this remarkable evolutionary radiation-that some trait constellations are viable and successful whereas others are not.Empirical support for a strongly limited set of viable trait combinations has accumulated for traits associated with single plant organs, such as leaves 7,9-12 , stems 13,14 and seeds [15][16][17] . Evidence across plant organs has been rarer, restricted geographically or taxonomically, and often contradictory [18][19][20][21][22][23][24][25][26][27][28][29] . How tightly whole-plant form and function are restricted at the global scale remains unresolved.Here we present the first global quantitative picture of essential functional diversity of extant vascular plants. We quantify the volume, shape and boundaries of this functional space via joint consideration of six traits that together capture the essence of plant form and function: adult plant height, stem specific density, leaf size expressed as leaf area, leaf mass per area, leaf nitrogen content per unit mass, and diaspore mass. Our dataset, based on a recently updated communal plant trait database 30 , covers 46,085 vascular plant species from 423 families and to our knowledge spans the widest range of growth-forms and geographical locations to date in published trait analyses, including some of the most extreme plant trait values ever measured in the field (Table 1, Extended Data Fig. 1). On this basis we reveal that the trait space actually occupied is strongly restricted as compared to four alternative null hypotheses. We demonstrate that plant species largely occupy a plane in the six-dimensional trait space. Two key trait dimensions within this plane are the size of whole plants and organs on the one hand, and the construction costs for photosynthetic leaf area, on the other. We subsequently show which sections of the plane are occupied, and how densely, by different growth-forms and major taxonomic groups. The design opportunities and limits indicated by today's global spectrum of plant form and function provide a foundation to achieve a better understanding of the evolutionary trajectory of vascular plants and help frame and test hypotheses as to where and Earth is home to a remarkable diversity of plant forms and life histories, yet comparatively few essential trait combinations have proved evolutionarily viable in today's terrestrial biosphere. By analysing worldwide variation in six major traits critical to growth, survival and reproduction within the largest sample of vascular plant species ever compiled, we found that occupancy of six-dimensional trait space is strongly concentrated, indicating coordination and trade-offs. Threequarters of trait variation is captured in a t...