SummaryRatios of nitrogen (N) isotopes in leaves could elucidate underlying patterns of N cycling across ecological gradients. To better understand global-scale patterns of N cycling, we compiled data on foliar N isotope ratios (δ 15 N), foliar N concentrations, mycorrhizal type and climate for over 11 000 plants worldwide. Arbuscular mycorrhizal, ectomycorrhizal, and ericoid mycorrhizal plants were depleted in foliar δ 15 N by 2‰, 3.2‰, 5.9‰, respectively, relative to nonmycorrhizal plants. Foliar δ 15 N increased with decreasing mean annual precipitation and with increasing mean annual temperature (MAT) across sites with MAT ≥ −0.5°C, but was invariant with MAT across sites with MAT < −0.5°C. In independent landscape-level to regionallevel studies, foliar δ 15 N increased with increasing N availability; at the global scale, foliar δ 15 N increased with increasing foliar N concentrations and decreasing foliar phosphorus (P) concentrations. Together, these results suggest that warm, dry ecosystems have the highest N availability, while plants with high N concentrations, on average, occupy sites with higher N availability than plants with low N concentrations. Global-scale comparisons of other components of the N cycle are still required for better mechanistic understanding of the determinants of variation in foliar δ 15 N and ultimately global patterns in N cycling.

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

Phosphorus (P) is generally considered the most common limiting nutrient for productivity of mature tropical lowland forests growing on highly weathered soils. It is often assumed that P limitation also applies to young tropical forests, but nitrogen (N) losses during land-use change may alter the stoichiometric balance of nutrient cycling processes. In the Amazon basin, about 16% of the original forest area has been cleared, and about 30-50% of cleared land is estimated now to be in some stage of secondary forest succession following agricultural abandonment. Here we use forest age chronosequences to demonstrate that young successional forests growing after agricultural abandonment on highly weathered lowland tropical soils exhibit conservative N-cycling properties much like those of N-limited forests on younger soils in temperate latitudes. As secondary succession progresses, N-cycling properties recover and the dominance of a conservative P cycle typical of mature lowland tropical forests re-emerges. These successional shifts in N:P cycling ratios with forest age provide a mechanistic explanation for initially lower and then gradually increasing soil emissions of the greenhouse gas nitrous oxide (N(2)O). The patterns of N and P cycling during secondary forest succession, demonstrated here over decadal timescales, are similar to N- and P-cycling patterns during primary succession as soils age over thousands and millions of years, thus revealing that N availability in terrestrial ecosystems is ephemeral and can be disrupted by either natural or anthropogenic disturbances at several timescales.

We analysed 1040 individual trees, located in 62 plots across the Amazon Basin for leaf mass per unit area (M-A), foliar carbon isotopic composition (delta C-13) 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 M-A, [C], [N] and [Mg] emerge as highly constrained by the taxonomic affiliation of tree species, but with others such as [P], [K], [Ca] and delta C-13 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 M-A and have a higher intrinsic [N], [P], [K], [Mg] and delta C-13 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 delta C-13 and reducing M-A. Mean annual temperature was negatively associated with leaf level [N], [P] and [K] concentrations. Total annual precipitation positively influences M-A, [C] and delta C-13, 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

Abstract. We analysed 1040 individual trees, positioned in sixty three plots across the Amazon Basin for leaf mass per area (MA), leaf carbon isotope composition (δ13C) and leaf level concentrations of C, N, P, Ca, Mg, K and Al. All trees were identified to the species with the dataset containing 58 families, 236 genera and 508 species, distributed across a wide range of soil types and precipitation regimes. Some foliar characters 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 composition 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.

Abstract. Vertical profiles in leaf mass per unit leaf area (M A ), foliar 13 C composition (δ 13 C), 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 M A and [C] DW (the subscript denoting on a dry weight basis) with negative within-tree gradients observed for δ 13 C, [Mg] Significant differences in within-tree gradients between individuals were observed only for M A , δ 13 C 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, A max . A new model is presented which is in agreement with the above observations. The model predicts that trees characterised by a low upper canopy A max should have shallow, or even non-existent, within-canopy gradients in A max , with optimal intra-canopy gradients becoming sharper as a tree's Published by Copernicus Publications on behalf of the European Geosciences Union. upper canopy A max increases. Nevertheless, in all cases it is predicted that the optimal within-canopy gradient in A max 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.

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Quantifying global patterns of terrestrial nitrogen (N) cycling is central to predicting future patterns of primary productivity, carbon sequestration, nutrient fluxes to aquatic systems, and climate forcing. With limited direct measures of soil N cycling at the global scale, syntheses of the 15N:14N ratio of soil organic matter across climate gradients provide key insights into understanding global patterns of N cycling. In synthesizing data from over 6000 soil samples, we show strong global relationships among soil N isotopes, mean annual temperature (MAT), mean annual precipitation (MAP), and the concentrations of organic carbon and clay in soil. In both hot ecosystems and dry ecosystems, soil organic matter was more enriched in 15N than in corresponding cold ecosystems or wet ecosystems. Below a MAT of 9.8°C, soil δ15N was invariant with MAT. At the global scale, soil organic C concentrations also declined with increasing MAT and decreasing MAP. After standardizing for variation among mineral soils in soil C and clay concentrations, soil δ15N showed no consistent trends across global climate and latitudinal gradients. Our analyses could place new constraints on interpretations of patterns of ecosystem N cycling and global budgets of gaseous N loss.