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Forest structure and dynamics vary across the Amazon Basin in an east-west gradient coincident with variations in soil fertility and geology. This has resulted in the hypothesis that soil fertility may play an important role in explaining Basin-wide variations in forest biomass, growth and stem turnover rates. Soil samples were collected in a total of 59 different forest plots across the Amazon Basin and analysed for exchangeable cations, carbon, nitrogen and pH, with several phosphorus fractions of likely different plant availability also quantified. Physical properties were additionally examined and an index of soil physical quality developed. Bivariate relationships of soil and climatic properties with above-ground wood productivity, stand-level tree turnover rates, above-ground wood biomass and wood density were first examined with multivariate regression models then applied. Both forms of analysis were undertaken with and without considerations regarding the underlying spatial structure of the dataset. Despite the presence of autocorrelated spatial structures complicating many analyses, forest structure and dynamics were found to be strongly and quantitatively related to edaphic as well as climatic conditions. Basin-wide differences in stand-level turnover rates are mostly influenced by soil physical properties with variations in rates of coarse wood production mostly related to soil phosphorus status. Total soil P was a better predictor of wood production rates than any of the fractionated organic- or inorganic-P pools. This suggests that it is not only the immediately available P forms, but probably the entire soil phosphorus pool that is interacting with forest growth on longer timescales. A role for soil potassium in modulating Amazon forest dynamics through its effects on stand-level wood density was also detected. Taking this into account, otherwise enigmatic variations in stand-level biomass across the Basin were then accounted for through the interacting effects of soil physical and chemical properties with climate. A hypothesis of self-maintaining forest dynamic feedback mechanisms initiated by edaphic conditions is proposed. It is further suggested that this is a major factor determining endogenous disturbance levels, species composition, and forest productivity across the Amazon Basin.

Many plant traits covary in a non-random manner reflecting interdependencies associated with "ecological strategy" dimensions. To understand how plants integrate their structural and physiological investments, data on leaf and leaflet size and the ratio of leaf area to sapwood area (ΦLS) obtained for 1020 individual trees (encompassing 661 species) located in 52 tropical forest plots across the Amazon Basin were incorporated into an analysis utilising existing data on species maximum height (Hmax), seed size, leaf mass per unit area (MA), foliar nutrients and δ13C, and branch xylem density (ρx). Utilising a common principal components approach allowing eigenvalues to vary between two soil fertility dependent species groups, five taxonomically controlled trait dimensions were identified. The first involves primarily cations, foliar carbon and MA and is associated with differences in foliar construction costs. The second relates to some components of the classic "leaf economic spectrum", but with increased individual leaf areas and a higher ΦLS newly identified components for tropical tree species. The third relates primarily to increasing Hmax and hence variations in light acquisition strategy involving greater MA, reductions in ΦLS and less negative δ13C. Although these first three dimensions were more important for species from high fertility sites the final two dimensions were more important for low fertility species and were associated with variations linked to reproductive and shade tolerance strategies. Environmental conditions influenced structural traits with ρx of individual species decreasing with increased soil fertility and higher temperatures. This soil fertility response appears to be synchronised with increases in foliar nutrient concentrations and reductions in foliar [C]. Leaf and leaflet area and ΦLS were less responsive to the environment than ρx. Thus, although genetically determined foliar traits such as those associated with leaf construction costs coordinate independently of structural characteristics such as maximum height, others such as the classical "leaf economic spectrum" covary with structural traits such as leaf size and ΦLS. Coordinated structural and physiological adaptions are also associated with light acquisition/shade tolerance strategies with several traits such as MA and [C] being significant components of more than one ecological strategy dimension. This is argued to be a consequence of a range of different potential underlying causes for any observed variation in such "ambiguous" traits. Environmental effects on structural and physiological characteristics are also coordinated but in a different way to the gamut of linkages associated with genotypic differences.

Tropical tree height-diameter (H:D) relationships may vary by forest type and region making large-scale estimates of above-ground biomass subject to bias if they ignore these differences in stem allometry. We have therefore developed a new global tropical forest database consisting of 39 955 concurrent H and D measurements encompassing 283 sites in 22 tropical countries. Utilising this database, our objectives were: 1. to determine if H:D relationships differ by geographic region and forest type (wet to dry forests, including zones of tension where forest and savanna overlap). 2. to ascertain if the H:D relationship is modulated by climate and/or forest structural characteristics (e.g. stand-level basal area, A). 3. to develop H:D allometric equations and evaluate biases to reduce error in future local-to-global estimates of tropical forest biomass. Annual precipitation coefficient of variation (PV), dry season length (SD), and mean annual air temperature (TA) emerged as key drivers of variation in H:D relationships at the pantropical and region scales. Vegetation structure also played a role with trees in forests of a high A being, on average, taller at any given D. After the effects of environment and forest structure are taken into account, two main regional groups can be identified. Forests in Asia, Africa and the Guyana Shield all have, on average, similar H:D relationships, but with trees in the forests of much of the Amazon Basin and tropical Australia typically being shorter at any given D than their counterparts elsewhere. The region-environment-structure model with the lowest Akaike's information criterion and lowest deviation estimated stand-level H across all plots to within amedian −2.7 to 0.9% of the true value. Some of the plot-to-plot variability in H:D relationships not accounted for by this model could be attributed to variations in soil physical conditions. Other things being equal, trees tend to be more slender in the absence of soil physical constraints, especially at smaller D. Pantropical and continental-level models provided less robust estimates of H, especially when the roles of climate and stand structure in modulating H:D allometry were not simultaneously taken into account.

No abstract available.

Vertical profiles in leaf mass per unit leaf area (MA), foliar 13C composition (δ13C), nitrogen (N), phosphorus (P), carbon (C) and major cation concentrations were estimated for 204 rain forest trees growing in 57 sites across the Amazon Basin. Data was analysed using a multilevel modelling approach, allowing a separation of gradients within individual tree canopies (within-tree gradients) as opposed to stand level gradients occurring because of systematic differences occurring between different trees of different heights (between-tree gradients). Significant positive within-tree gradients (i.e. increasing values with increasing sampling height) were observed for MA and [C]DW (the subscript denoting on a dry weight basis) with negative within-tree gradients observed for δ13C, [Mg]DW and [K]DW. No significant within-tree gradients were observed for [N]DW, [P]DW or [Ca]DW. The magnitudes of between-tree gradients were not significantly different to the within-tree gradients for MA, δ13C, [C]DW, [K]DW, [N]DW, [P]DW and [Ca]DW. But for [Mg]DW, although there was no systematic difference observed between trees of different heights, strongly negative within-tree gradients were found to occur. When expressed on a leaf area basis (denoted by the subscript "A"), significant positive gradients were observed for [N]A, [P]A and [K]A both within and between trees, these being attributable to the positive intra- and between-tree gradients in MA mentioned above. No systematic within-tree gradient was observed for either [Ca]A or [Mg]A, but with a significant positive gradient observed for [Mg]A between trees (i.e. with taller trees tending to have a higher Mg per unit leaf area). Significant differences in within-tree gradients between individuals were observed only for MA, δ13C and [P] A. This was best associated with the overall average [P]A for each tree, this also being considered to be a surrogate for a tree's average leaf area based photosynthetic capacity, Amax. A new model is presented which is in agreement with the above observations. The model predicts that trees characterised by a low upper canopy Amax should have shallow, or even non-existent, within-canopy gradients in Amax, with optimal intra-canopy gradients becoming sharper as a tree's upper canopy Amax increases. Nevertheless, in all cases it is predicted that the optimal within-canopy gradient in Amax should be substantially less than for photon irradiance. Although this is also shown to be consistent with numerous observations as illustrated by a literature survey of gradients in photosynthetic capacity for broadleaf trees, it is also in contrast to previously held notions of optimality. A new equation relating gradients in photosynthetic capacity within broadleaf tree canopies to the photosynthetic capacity of their upper canopy leaves is presented.

Soil samples were collected in six South American countries in a total of 71 different 1 ha forest plots across the Amazon Basin as part of the RAINFOR project. They were analysed for total and exchangeable cations, C, N, pH with various P fractions also determined. Physical properties were also examined and an index of soil physical quality proposed. A diverse range of soils was found. For the western areas near the Andean cordillera and the southern and northern fringes, soils tend to be distributed among the lower pedogenetic levels, while the central and eastern areas of Amazonia have more intensely weathered soils. This gives rise to a large variation of soil chemical and physical properties across the Basin, with soil properties varying predictably along a gradient of pedogenic development. Nutrient pools generally increased slightly in concentration from the youngest to the intermediate aged soils after which a gradual decline was observed with the lowest values found in the most weathered soils. Soil physical properties were strongly correlated with soil fertility, with favourable physical properties occurring in highly weathered and nutrient depleted soils and with the least weathered, more fertile soils having higher incidence of limiting physical properties. Soil phosphorus concentrations varied markedly in accordance with weathering extent and appear to exert an important influence on the nitrogen cycle of Amazon forest soils.

The production of aboveground soft tissue represents an important share of total net primary production in tropical rain forests. Here we draw from a large number of published and unpublished datasets (n=81 sites) to assess the determinants of litterfall variation across South American tropical forests. We show that across old-growth tropical rainforests, litterfall averages 8.61±1.91 Mg ha−1 yr−1 (mean ± standard deviation, in dry mass units). Secondary forests have a lower annual litterfall than old-growth tropical forests with a mean of 8.01±3.41 Mg ha−1 yr−1. Annual litterfall shows no significant variation with total annual rainfall, either globally or within forest types. It does not vary consistently with soil type, except in the poorest soils (white sand soils), where litterfall is significantly lower than in other soil types (5.42±1.91 Mg ha−1 yr−1). We also study the determinants of litterfall seasonality, and find that it does not depend on annual rainfall or on soil type. However, litterfall seasonality is significantly positively correlated with rainfall seasonality. Finally, we assess how much carbon is stored in reproductive organs relative to photosynthetic organs. Mean leaf fall is 5.74±1.83 Mg ha−1 yr−1 (71% of total litterfall). Mean allocation into reproductive organs is 0.69±0.40 Mg ha−1 yr−1 (9% of total litterfall). The investment into reproductive organs divided by leaf litterfall increases with soil fertility, suggesting that on poor soils, the allocation to photosynthetic organs is prioritized over that to reproduction. Finally, we discuss the ecological and biogeochemical implications of these results.

It has been hypothesized that as soil fertility increases, the amount of carbon allocated to below-ground production (fine roots) should decrease. To evaluate this hypothesis, we measured the standing crop fine root mass and the production of fine roots (−1 yr−1, method 1 and 2, respectively) as much as for the more fertile loamy soil forest (1.51±0.14, method 1, and from 1.03±0.31 to 1.36±0.23 Mg C ha−1 yr−1, method 2). Similarly, the average of standing crop fine root mass was higher in the white-sands forest (10.94±0.33 Mg C ha−1) as compared to the forest on the more fertile soil (from 3.04±0.15 to 3.64±0.18 Mg C ha−1). The standing crop fine root mass also showed a temporal pattern related to rainfall, with the production of fine roots decreasing substantially in the dry period of the year 2005. These results suggest that soil resources may play an important role in patterns of carbon allocation to the production of fine roots in these forests as the proportion of carbon allocated to above- and below-ground organs is different between forest types. Thus, a trade-off between above- and below-ground growth seems to exist with our results also suggesting that there are no differences in total net primary productivity between these two forests, but with higher below-ground production and lower above-ground production for the forest on the nutrient poor soil.

The net primary productivity (NPP) of tropical forests is one of the most important and least quantified components of the global carbon cycle. Most relevant studies have focused particularly on the quantification of the above-ground coarse wood productivity, and little is known about the carbon fluxes involved in other elements of the NPP, the partitioning of total NPP between its above- and below-ground components and the main environmental drivers of these patterns. In this study we quantify the above- and below-ground NPP of ten Amazonian forests to address two questions: (1) How do Amazonian forests allocate productivity among its above- and below-ground components? (2) How do soil and leaf nutrient status and soil texture affect the productivity of Amazonian forests? Using a standardized methodology to measure the major elements of productivity, we show that NPP varies between 9.3±1.3 Mg C ha−1 yr−1 (mean±standard error), at a white sand plot, and 17.0±1.4 Mg C ha−1 yr−1 at a very fertile Terra Preta site, with an overall average of 12.8±0.9 Mg C ha−1 yr−1. The studied forests allocate on average 64±3% and 36±3% of the total NPP to the above- and below-ground components, respectively. The ratio of above-ground and below-ground NPP is almost invariant with total NPP. Litterfall and fine root production both increase with total NPP, while stem production shows no overall trend. Total NPP tends to increase with soil phosphorus and leaf nitrogen status. However, allocation of NPP to below-ground shows no relationship to soil fertility, but appears to decrease with the increase of soil clay content.

We analysed 1040 individual trees, located in 62 plots across the Amazon Basin for leaf mass per unit area (MA), foliar carbon isotopic composition (δ13C) and leaf level concentrations of C, N, P, Ca, Mg, K and Al. All trees were identified to the species level with the dataset containing 58 families, 236 genera and 508 species, distributed across a wide range of soil types and precipitation regimes. Some foliar characteristics such as MA, [C], [N] and [Mg] emerge as highly constrained by the taxonomic affiliation of tree species, but with others such as [P], [K], [Ca] and δ13C also strongly influenced by site growing conditions. By removing the environmental contribution to trait variation, we find that intrinsic values of most trait pairs coordinate, although different species (characterised by different trait suites) are found at discrete locations along a common axis of coordination. Species that tend to occupy higher fertility soils are characterised by a lower MA and have a higher intrinsic [N], [P], [K], [Mg] and δ13C than their lower fertility counterparts. Despite this consistency, different scaling patterns were observed between low and high fertility sites. Inter-relationships are thus substantially modified by growth environment. Analysing the environmental component of trait variation, we found soil fertility to be the most important predictor, influencing all leaf nutrient concentrations and δ13C and reducing MA. Mean annual temperature was negatively associated with leaf level [N], [P] and [K] concentrations. Total annual precipitation positively influences MA, [C] and δ13C, but with a negative impact on [Mg]. These results provide a first basis for understanding the relationship between the physiological functioning and distribution of tree species across Amazonia.

Given the importance of Amazon rainforest in the global carbon and hydrological cycles, there is a need to parameterize and validate ecosystem gas exchange and vegetation models for this region in order to adequately simulate present and future carbon and water balances. In this study, a sun and shade canopy gas exchange model is calibrated and evaluated at five rainforest sites using eddy correlation measurements of carbon and energy fluxes. Results from the model-data evaluation suggest that with adequate parameterisation, photosynthesis models taking into account the separation of diffuse and direct irradiance and the dynamics of sunlit and shaded leaves can accurately represent photosynthesis in these forests. Also, stomatal conductance formulations that only take into account atmospheric demand fail to correctly simulate moisture and CO2 fluxes in forests with a pronounced dry season, particularly during afternoon conditions. Nevertheless, it is also the case that large uncertainties are associated not only with the eddy correlation data, but also with the estimates of ecosystem respiration required for model validation. To accurately simulate Gross Primary Productivity (GPP) and energy partitioning the most critical parameters and model processes are the quantum yield of photosynthetic uptake, the maximum carboxylation capacity of Rubisco, and simulation of stomatal conductance. Using this model-data synergy, we developed scaling functions to provide estimates of canopy photosynthetic parameters for a range of diverse forests across the Amazon region, utilising the best fitted parameter for maximum carboxylation capacity of Rubisco, and foliar nutrients (N and P) for all sites.

Xylem density is a physical property of wood that varies between individuals, species and environments. It reflects the physiological strategies of trees that lead to growth, survival and reproduction. Measurements of branch xylem density, ρx, were made for 1653 trees representing 598 species, sampled from 87 sites across the Amazon basin. Measured values ranged from 218 kg m−3 for a Cordia sagotii (Boraginaceae) from Mountagne de Tortue, French Guiana to 1130 kg m−3 for an Aiouea sp. (Lauraceae) from Caxiuana, Central Pará, Brazil. Analysis of variance showed significant differences in average ρx across regions and sampled plots as well as significant differences between families, genera and species. A partitioning of the total variance in the dataset showed that species identity (family, genera and species) accounted for 33% with environment (geographic location and plot) accounting for an additional 26%; the remaining "residual" variance accounted for 41% of the total variance. Variations in plot means, were, however, not only accountable by differences in species composition because xylem density of the most widely distributed species in our dataset varied systematically from plot to plot. Thus, as well as having a genetic component, branch xylem density is a plastic trait that, for any given species, varies according to where the tree is growing in a predictable manner. Within the analysed taxa, exceptions to this general rule seem to be pioneer species belonging for example to the Urticaceae whose branch xylem density is more constrained than most species sampled in this study. These patterns of variation of branch xylem density across Amazonia suggest a large functional diversity amongst Amazonian trees which is not well understood.