Terrestrial carbon stock mapping is important for the successful implementation of climate change mitigation policies. Its accuracy depends on the availability of reliable allometric models to infer oven-dry aboveground biomass of trees from census data. The degree of uncertainty associated with previously published pantropical aboveground biomass allometries is large. We analyzed a global database of directly harvested trees at 58 sites, spanning a wide range of climatic conditions and vegetation types (4004 trees ≥ 5 cm trunk diameter). When trunk diameter, total tree height, and wood specific gravity were included in the aboveground biomass model as covariates, a single model was found to hold across tropical vegetation types, with no detectable effect of region or environmental factors. The mean percent bias and variance of this model was only slightly higher than that of locally fitted models. Wood specific gravity was an important predictor of aboveground biomass, especially when including a much broader range of vegetation types than previous studies. The generic tree diameter-height relationship depended linearly on a bioclimatic stress variable E, which compounds indices of temperature variability, precipitation variability, and drought intensity. For cases in which total tree height is unavailable for aboveground biomass estimation, a pantropical model incorporating wood density, trunk diameter, and the variable E outperformed previously published models without height. However, to minimize bias, the development of locally derived diameter-height relationships is advised whenever possible. Both new allometric models should contribute to improve the accuracy of biomass assessment protocols in tropical vegetation types, and to advancing our understanding of architectural and evolutionary constraints on woody plant development.

Summary 1.The 'functional traits' of species have been heralded as promising predictors for species' demographic rates and life history. Multiple studies have linked plant species' demographic rates to commonly measured traits. However, predictive power is usually low -raising questions about the practical usefulness of traits -and analyses have been limited to size-independent univariate approaches restricted to a particular life stage. 2. Here we directly evaluated the predictive power of multiple traits simultaneously across the entire life cycle of 136 tropical tree species from central Panama. Using a model-averaging approach, we related wood density, seed mass, leaf mass per area and adult stature (maximum diameter) to onset of reproduction, seed production, seedling establishment, and growth and survival at seedling, sapling and adult stages. 3. Three of the four traits analysed here (wood density, seed mass and adult stature) typically explained 20-60% of interspecific variation at a given vital rate and life stage. There were strong shifts in the importance of different traits throughout the life cycle of trees, with seed mass and adult stature being most important early in life, and wood density becoming most important after establishment. Every trait had opposing effects on different vital rates or at different life stages; for example, seed mass was associated with higher seedling establishment and lower initial survival, wood density with higher survival and lower growth, and adult stature with decreased juvenile but increased adult growth and survival. 4. Forest dynamics are driven by the combined effects of all demographic processes across the full life cycle. Application of a multitrait and full-life cycle approach revealed the full role of key traits, and illuminated how trait effects on demography change through the life cycle. The effects of traits on one life stage or vital rate were sometimes offset by opposing effects at another stage, revealing the danger of drawing broad conclusions about functional traitdemography relationships from analysis of a single life stage or vital rate. Robust ecological and evolutionary conclusions about the roles of functional traits rely on an understanding of the relationships of traits to vital rates across all life stages.

In this paper, an outline of the methods for measuring seed limitation, establishment limitation and their components are presented. These methods are applicable to any study that quantifies seed rain at an unbiased sample of locations in a community or explicitly measures the shapes of seed shadows. The usefulness of these methods were evaluated by using them in several species in tropical forest. The implications of observed seed and establishment limitation for tropical forest diversity and conservation are assessed.

Patterns of size inequality in crowded plant populations are often taken to be indicative of the degree of size asymmetry of competition, but recent research suggests that some of the patterns attributed to size-asymmetric competition could be due to spatial structure. To investigate the theoretical relationships between plant density, spatial pattern, and competitive size asymmetry in determining size variation in crowded plant populations, we developed a spatially explicit, individual-based plant competition model based on overlapping zones of influence. The zone of influence of each plant is modeled as a circle, growing in two dimensions, and is allometrically related to plant biomass. The area of the circle represents resources potentially available to the plant, and plants compete for resources in areas in which they overlap. The size asymmetry of competition is reflected in the rules for dividing up the overlapping areas. Theoretical plant populations were grown in random and in perfectly uniform spatial patterns at four densities under size-asymmetric and size-symmetric competition. Both spatial pattern and size asymmetry contributed to size variation, but their relative importance varied greatly over density and over time. Early in stand development, spatial pattern was more important than the symmetry of competition in determining the degree of size variation within the population, but after plants grew and competition intensified, the size asymmetry of competition became a much more important * E-mail: jw@kvl.dk. † E-mail: stoll@sgi.unibe.ch. ‡ E-mail: helene@eno.princeton.edu. § E-mail: ajasen@concentric.net. source of size variation. Size variability was slightly higher at higher densities when competition was symmetric and plants were distributed nonuniformly in space. In a uniform spatial pattern, size variation increased with density only when competition was size asymmetric. Our results suggest that when competition is size asymmetric and intense, it will be more important in generating size variation than is local variation in density. Our results and the available data are consistent with the hypothesis that high levels of size inequality commonly observed within crowded plant populations are largely due to size-asymmetric competition, not to variation in local density.Keywords: asymmetric competition, individual-based models, population structure, size inequality, spatial effects, zone of influence.Competition among individuals usually increases size variation within plant populations, but there is controversy over the mechanisms through which this occurs. This controversy reflects a fundamental disagreement about the nature of competition among individual plants. Some studies have concluded that a major factor generating size variation in crowded plant populations is the "size asymmetry" of competition: larger plants have a disproportionate advantage (for their relative size) in competition with smaller plants, suppressing their growth (Begon 1984;Weiner 1990;Schwinning and Weiner 1...

Abstract. Numerous grassland experiments have found evidence for a complementarity effect, an increase in productivity with higher plant species richness due to niche partitioning. However, empirical tests of complementarity in natural forests are rare. We conducted a spatially explicit analysis of 518 433 growth records for 274 species from a 50-ha tropical forest plot to test neighborhood complementarity, the idea that a tree grows faster when it is surrounded by more dissimilar neighbors. We found evidence for complementarity: focal tree growth rates increased by 39.8% and 34.2% with a doubling of neighborhood multi-trait dissimilarity and phylogenetic dissimilarity, respectively. Dissimilarity from neighbors in maximum height had the most important effect on tree growth among the six traits examined, and indeed, its effect trended much larger than that of the multitrait dissimilarity index. Neighborhood complementarity effects were strongest for lightdemanding species, and decreased in importance with increasing shade tolerance of the focal individuals. Simulations demonstrated that the observed neighborhood complementarities were sufficient to produce positive stand-level biodiversity-productivity relationships. We conclude that neighborhood complementarity is important for productivity in this tropical forest, and that scaling down to individual-level processes can advance our understanding of the mechanisms underlying stand-level biodiversity-productivity relationships.