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 ecosystem-level 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 traits, leaf traits, root and stem traits and regenerative traits, and puts particular emphasis on traits important for predicting species’ effects on key ecosystem properties. We hope this new handbook becomes a standard companion in local and global efforts to learn about the responses and impacts of different plant species with respect to environmental changes in the present, past and future.

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New handbook for standardised measurement of plant functional traits worldwide

Fire is a worldwide phenomenon that appears in the geological record soon after the appearance of terrestrial plants. Fire influences global ecosystem patterns and processes, including vegetation distribution and structure, the carbon cycle, and climate. Although humans and fire have always coexisted, our capacity to manage fire remains imperfect and may become more difficult in the future as climate change alters fire regimes. This risk is difficult to assess, however, because fires are still poorly represented in global models. Here, we discuss some of the most important issues involved in developing a better understanding of the role of fire in the Earth system.

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Fire in the Earth System

Two features characterize all natural communities. First, they are dynamic systems. The densities and age-structures of populations change with time, as do the relative abundances of species; local extinctions are commonplace (37). For many communities, a self-reproducing climax state may only exist as an average condition on a relatively large spatial scale, and even that has yet to be rigorously demonstrated (36). The idea that equilibrium is rarely achieved on the local scale was expressed decades ago by a number of forest ecologists (e.g. 10 1, 168). One might even argue that continued application of the concept of climax to natural systems is simply an exercise in metaphysics (41). While this view may seem extreme, major climatic shifts often recur at time intervals shorter than that required for a community to reach competitive equilibrium or alter the geographical distributions of species (6, 21, 43, 76, 92). Climatic variation of this kind influences ecological patterns over large areas, sometimes encompassing entire continents. Other agents of temporal change in natural communities operate over a wide range of smaller spatial scales (47, 242). Second, natural communities are spatially heterogeneous. This statement is true at any scale of resolution (242), but it is especially apparent on what is commonly referred to as the regional scale. (By region I mean an area that potentially encompasses more than one colonizable patch.) Across any land or seascape, one observes a mosaic of patches identified by spatial discontinuities in the distributions of populations (153, 159, 161, 231, 239, 240). Closer examination often reveals a smaller-scale patchwork of same-aged individuals (e.g. 85-87, 101, 146, 199,204,217-220,235,246). Discrete patch boundaries sometimes reflect species-specific responses to

The Role of Disturbance in Natural Communities

We discuss the dynamics of plant litter, the effects of litter on the chemical and physical environment, the direct and indirect effects of plant litter on plant populations and communities, and different adaptative traits that may be related to litter accumulation. The production of litter depends primarily on the site productivity, but other properties of the environment, as well as chance, may introduce important variation. The existence of time lags between the production of plant organs and their transformation into litter appears as a relevant character of litter dynamics seldom included in models. Herbivory, and other processes that destroy biomass or reduce productivity, may reduce the amount of litter produced. The destruction of litter encompasses a complex of interactions. The main processes, including physical and chemical degradation, consumption by invertebrates and decomposition, are differentially affected by the environment and by the physical and chemical characteristics of the litter itself. The relative importance of those processes varies among systems.

Plant litter : its dynamics and effects on plant community structure

In order to estimate the production of charcoal and the atmospheric emissions of trace gases volatilized by burning we have estimated the global amounts of biomass which are affected by fires. We have roughly calculated annual gross burning rates ranging between about 5 Pg and 9 Pg (1 Pg = 1015 g) of dry matter (2–4 Pg C). In comparison, about 9–17 Pg of above-ground dry matter (4–8 Pg C) is exposed to fires, indicating a worldwide average burning efficiency of about 50%. The production of dead below-ground dry matter varies between 6–9 Pg per year. We have tentatively indicated the possibility of a large production of elemental carbon (0.5–1.7 Pg C/yr) due to the incomplete combustion of biomass to charcoal. This provides a sink for atmospheric CO2, which would have been particularly important during the past centuries. From meager statistical information and often ill-documented statements in the literature, it is extremely difficult to calculate the net carbon release rates to the atmosphere from the biomass changes which take place, especially in the tropics. All together, we calculate an overall effect lof the biosphere on the atmospheric carbon dioxide budget which may range between the possibilities of a net uptake or a net release of about 2 Pg C/yr. The release of CO2 to the atmosphere by deforestation projects may well be balanced by reforestation and by the production of charcoal. Better information is needed, however, to make these estimates more reliable.

Estimates of gross and net fluxes of carbon between the biosphere and the atmosphere from biomass burning

Plant species which have survived fires for tens of thousands of years may not only have selected survival mechanisms, but also inherent flammable properties that contribute to the perpetuation of fire—dependent plant communities. This concept goes by beyond the commonly accepted fire climate—fuel moisture basis of wildland fire occurrence. Plant communities may be ignited accidentally or randomly, but the character of burning is not random. The following hypothesis treats this interaction between fire and the ecosystem: Fire—dependent plant communities burn more readily than non—fire—dependent communities because natural selection has favored development of characteristics that make them more flammable. The hypothesis was experimentally derived following laboratory combustion tests with litter of eucalyptus (Eucalyptus obliqua L'Herit), ponderosa pine (Pinus ponderosa Laws.), and tropical hardwood leaves.

Robert W. Mutch

Papers

Wildland Fires and Ecosystems--A Hypothesis