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 ...

Worldwide decomposition rates depend both on climate and the legacy of plant functional traits as litter quality. To quantify the degree to which functional differentiation among species affects their litter decomposition rates, we brought together leaf trait and litter mass loss data for 818 species from 66 decomposition experiments on six continents. We show that: (i) the magnitude of species-driven differences is much larger than previously thought and greater than climate-driven variation; (ii) the decomposability of a species' litter is consistently correlated with that species' ecological strategy within different ecosystems globally, representing a new connection between whole plant carbon strategy and biogeochemical cycling. This connection between plant strategies and decomposability is crucial for both understanding vegetation-soil feedbacks, and for improving forecasts of the global carbon cycle.

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 ...

Associations between specific leaf area (SLA), leaf water content (LWC) and leaf thickness (LT) in 77 species were analysed to identify which of these traits gave a better indicator value of general plant resource-use strategy within the flora of central-western Argentina, in which succulent species are common. • When all species were considered together, SLA and LWC were not significantly correlated. All high-SLA tender-leafed species showed high LWC. Low SLA, however, was associated both with low LWC (sclerophyllous species) and with high LWC (succulents). When succulents were excluded, the association between SLA and LWC was significant and positive. A similar trend was found for a mixed set of nonsucculent species from other floras of the world. • In the Argentine data set, SLA and LT, but not LWC, were significantly correlated with species' scores along a multivariate axis of plant resource-use strategy. • Because of its clearer ecological interpretation and its applicability across different floras, SLA appears to be the best candidate for inclusion in large comparative databases.

Aim  To determine how the distribution and cover of different vegetation types are affected by physical factors and livestock in a mountain range with a long evolutionary history of grazing. Location  Upper vegetation belt of the Córdoba mountains (1700–2800 m a.s.l., 31º34′ S, 64º50′ W) in central Argentina. Methods  Using GIS, we analysed the relationships of plant cover types to physical features (physiography and topography) and indicators of accumulated livestock pressure (distance to human settlements and roads) through multinomial logistic regression. We predicted a present vegetation map which was validated with a real map. We then constructed two maps simulating minimum and maximum values of accumulated livestock pressure for the whole area. Map comparisons allowed evaluation of the possible influence of livestock, both in extension and intensity. Results  Both physical features and livestock pressure influenced the occurrence of vegetation units. The overall accuracy of the predicted map at the pixel level was low (26%) indicating low habitat specificity of the vegetation units. We suggest that some part of the unaccounted for variance was due to livestock pressure patterns that were not fully captured by our indicators. Our models proved adequate for predicting the total percentages of vegetation units at coarser scales. The extrapolations showed that under a history of low livestock pressure, such as in sites far away from human settlements and roads, the area would be dominated by woodlands, tussock grasslands and natural rock outcrops. Under a history of heavy livestock pressure, in turn, rock exposed by erosion, tussock grasslands and natural rock outcrops would dominate. Main conclusions  Vegetation units showed low habitat specificity, and were associated with accumulated livestock pressure, indicating that livestock and its associated activities are important factors structuring the landscape and have important consequences for the integrity of the ecosystem. Results suggest that although this system evolved with large herbivores, it has experienced irreversible degradation processes, and intensification of current domestic livestock pressure is likely to lead to even more land degradation.

Despite the vast diversity and complexity of herbivores, plants and their interactions, most authors agree that a small number of components of leaf quality affect preference by generalist herbivores in a predictable way. However, herbivore preference is determined not only by intrinsic plant attributes and herbivore biology but also by the environmental context. Within this framework, we aimed to analyse general interspecific trends in the association between herbivory and leaf traits over a wide range of angiosperms from central Argentina. We (i) tested for consistent associations between leaf traits, consumption in the field, and preference of generalist invertebrate herbivores in cafeteria experiments; (ii) assessed how well herbivore preferences in cafeterias matched leaf consumption in the field; and (iii) developed a simple conceptual model linking leaf traits, herbivore preference in cafeterias and consumption in the field. In general, we found that tender leaves with higher nutritional quality were preferred by herbivores, both in the field and in cafeteria experiments. According to our model, this relationship between field and cafeteria consumption and leaf quality is observed when generalist herbivores and plants of high accessibility are considered. However, differences between leaf consumption in the field and in cafeteria experiments can also be found. At least two reasons can account for this: (i) specialized plant-herbivore relationships often occur in the field, whereas cafeteria experiments tend to consider only one or a few generalist herbivores; (ii) different plant species growing in the field often differ in their degree of accessibility to herbivores, whereas in cafeteria experiments all species are equally accessible. Our results add new evidence to a growing consensus that, although herbivory in the field is determined by many factors, consistent patterns of differential susceptibility to foliar feeders can be found in leaves differing in nutritional quality and thus in resource-use strategy.

Questions Most vegetation descriptions tacitly assume that floristic composition and physiognomy are tightly linked. However, the two vegetation properties may not respond in a similar way to environmental and disturbance gradients, leading to uninformed management planning and difficulties when attempting to restore degraded ecosystems. In this context, we addressed two main questions: (1) how close are relations between floristic and physiognomic types as defined by numerical vegetation classification in mountain ecosystems; and (2) how are floristic and physiognomic types distributed along the elevation gradient? Location Central mountains of Argentina, between 500 and 1700 m a.s.l. Methods We selected 437 sites where we performed complete floristic and physiognomic relevés. We classified eight physiognomic and eight floristic types. We tested the relationship between the two classifications through a chi square analysis. We tested the association between elevation and each physiognomic and floristic type with random permutations. Results In general, floristic types were significantly and positively associated with more than one physiognomic type and vice versa. Physiognomic and floristic types responded differently to the elevation gradient. Floristic types were restricted to different sections of the gradient, although having large overlap among them. In contrast, seven out of the eight physiognomic types did not show elevation restriction, being distributed along the complete elevation gradient. The open low woodland with shrubs was the only restricted physiognomy, significantly absent from the upper part of the gradient. Conclusions We highlight the importance of considering the two vegetation properties independently when characterizing vegetation patterns in heterogeneous systems, since they show decoupled responses to environmental gradients. We note that the assumption of a direct link between floristic composition and physiognomy may induce bias into the understanding of vegetation patterns and processes. Hence, we encourage managers and restoration practitioners to consider the complete range of possible physiognomic types under each floristic type.