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.

Abstract. Understanding secondary successional processes in Amazonian terrestrial ecosystems is becoming increasingly important as continued deforestation expands the area that has become secondary forest, or at least has been through a recent phase of secondary forest growth. Most Amazonian soils are highly weathered and relatively nutrient poor, but the role of nutrients as a factor determining successional processes is unclear. Soils testing and chronosequence studies have yielded equivocal results regarding the possible role of nutrient limitation. The objective of this paper is to report the first two years' results of a nitrogen (N) and phosphorus (P) fertilization experiment in a 6-yr-old secondary forest growing on an abandoned cattle pasture on a clayey Oxisol. Growth of remnant grasses responded significantly to the N ϩ P treatment, whereas tree biomass increased significantly following N-only and N ϩ P treatments. The plants took up about 10% of the 50 kg P/ha of the first year's application, and recovery in soil fractions could account for the rest. The trees took up about 20% of the 100 kg N/ha of the first year's application. No changes in soil inorganic N, soil microbial biomass N, or litter decomposition rates have been observed so far, but soil faunal abundances increased in fertilized plots relative to the control in the second year of the study. A pulse of nitric oxide and nitrous oxide emissions was measured in the N-treated plots only shortly after the second year's application. Net N mineralization and net nitrification assays demonstrated strong immobilization potential, indicating that much of the N was probably retained in the large soil organic-N pool. Although P availability is low in these soils and may partially limit biomass growth, the most striking result of this study so far is the significant response of tree growth to N fertilization. Repeated fire and other losses of N from degraded pastures may render tree growth N limited in some young Amazonian forests. Changes in species composition and monitoring of long-term effects on biomass accumulation will be addressed as this experiment is continued.

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[1] Changes in land-use and climate are likely to alter moisture and substrate availability in tropical forest soils, but quantitative assessment of the role of resource constraints as regulators of soil trace gas fluxes is rather limited. The primary objective of this study was to quantify the effects of moisture and substrate availability on soil trace gas fluxes in an Amazonian regrowth forest. We measured the efflux of carbon dioxide (CO 2 ), nitric oxide (NO), nitrous oxide (N 2 O), and methane (CH 4 ) from soil in response to two experimental manipulations. In the first, we increased soil moisture availability during the dry season by irrigation; in the second, we decreased substrate availability by continuous removal of aboveground litter. In the absence of irrigation, soil CO 2 efflux decreased during the dry season while irrigation maintained soil CO 2 efflux levels similar to the wet season. Large variations in soil CO 2 efflux consistent with a significant moisture constraint on respiration were observed in response to soil wet-up and dry-down events. Annual soil C efflux for irrigated plots was 27 and 13% higher than for control plots in 2001 and 2002, respectively. Litter removal significantly reduced soil CO 2 efflux; annual soil C efflux in 2002 was 28% lower for litter removal plots compared to control plots. The annual soil C efflux:litterfall C ratio for the control treatment (4.0-5.2) was consistent with previously reported values for regrowth forests that indicate a relatively large belowground C allocation. In general, fluxes of N 2 O and CH 4 were higher during the wet season and both fluxes increased during dry-season irrigation. There was no seasonal effect on NO fluxes. Litter removal had no significant impact on N oxide or CH 4 emissions. Net soil nitrification did not respond to dry-season irrigation, but was somewhat reduced by litter removal. Overall, these results demonstrate significant soil moisture and substrate constraints on soil trace gas emissions, particularly for CO 2 , and suggest that climate and land-use changes that alter moisture and substrate availability are therefore likely to have an impact on atmosphere chemistry.

Fires set for slash-and-burn agriculture contribute to the current unsustainable accumulation of atmospheric greenhouse gases, and they also deplete the soil of essential nutrients, which compromises agricultural sustainability at local scales. Integrated assessments of greenhouse gas emissions have compared intensive cropping systems in industrialized countries, but such assessments have not been applied to common cropping systems of smallholder farmers in developing countries. We report an integrated assessment of greenhouse gas emissions in slash-and-burn agriculture and an alternative chopand-mulch system in the Amazon Basin. The soil consumed atmospheric methane (CH 4 ) under slash-and-burn treatment and became a net emitter of CH 4 to the atmosphere under the mulch treatment. Mulching also caused about a 50% increase in soil emissions of nitric oxide and nitrous oxide and required greater use of fertilizer and fuel for farm machinery. Despite these significantly higher emissions of greenhouse gases during the cropping phase under the alternative chop-and-mulch system, calculated pyrogenic emissions in the slash-and-burn system were much larger, especially for CH 4 . The global warming potential CO 2 -equivalent emissions calculated for the entire crop cycles were at least five times lower in chop-and-mulch compared with slash-and-burn. The crop yields were similar for the two systems. While economic and logistical considerations remain to be worked out for alternatives to slash-and-burn, these results demonstrate a potential 'win-win' strategy for maintaining soil fertility and reducing net greenhouse gas emissions, thus simultaneously contributing to sustainability at both spatial scales.

The absorption of atmospheric water directly into leaves enables plants to alleviate the water stress caused by low soil moisture, hydraulic resistance in the xylem and the effect of gravity on the water column, while enabling plants to scavenge small inputs of water from leaf‐wetting events. By increasing the availability of water, and supplying it from the top of the canopy (in a direction facilitated by gravity), foliar uptake (FU) may be a significant process in determining how forests interact with climate, and could alter our interpretation of current metrics for hydraulic stress and sensitivity. FU has not been reported for lowland tropical rainforests; we test whether FU occurs in six common Amazonian tree genera in lowland Amazônia, and make a first estimation of its contribution to canopy–atmosphere water exchange. We demonstrate that FU occurs in all six genera and that dew‐derived water may therefore be used to “pay” for some morning transpiration in the dry season. Using meteorological and canopy wetness data, coupled with empirically derived estimates of leaf conductance to FU (kfu), we estimate that the contribution by FU to annual transpiration at this site has a median value of 8.2% (103 mm/year) and an interquartile range of 3.4%–15.3%, with the biggest sources of uncertainty being kfu and the proportion of time the canopy is wet. Our results indicate that FU is likely to be a common strategy and may have significant implications for the Amazon carbon budget. The process of foliar water uptake may also have a profound impact on the drought tolerance of individual Amazonian trees and tree species, and on the cycling of water and carbon, regionally and globally.

-The physiological and morphological responses of the forage grasses Brachiaria brizantha cv. Marandu, B. decumbens and B. humidicola were compared for plants grown in pots under flooding and well-drained conditions for 14 days. Flooding reduced specific leaf area and biomass allocation to roots in all species and enhanced leaf senescence in B. brizantha and B. decumbens. Relative growth rate was reduced by flooding in B. brizantha and B. decumbens, but not in B. humidicola. Leaf elongation rate was unaffected by flooding in B. decumbens and B. humidicola, but declined in B. brizantha since the first day of flooding. Net photosynthesis and leaf chlorophyll content were reduced by flooding in B. brizantha; however, no flooding effect could be detected in the other two species. For all species, there was a close relationship between net photosynthesis and stomatal conductance under flooding. These results show that the studied species have distinct degrees of tolerance to flood, B. brizantha is intolerant, B. decumbens is moderately tolerant and B. humidicola is tolerant. Because leaf elongation rate was immediately depressed by flooding only in B. brizantha, this measurement could be appropriate as an early detection mechanism for relative flood tolerance in Brachiaria spp.Index terms: Brachiaria brizantha, Brachiaria decumbens, Brachiaria humidicola, leaf area, chlorophyll, photosynthesis, biomass, growth rate.RESPOSTAS MORFOLÓGICAS E FISIOLÓGICAS DE BRACHIARIA SPP. AO ALAGAMENTO DO SOLO RESUMO -As respostas morfológicas e fisiológicas de Brachiaria brizantha cv. Marandu, B. decumbens e B. humidicola foram comparadas em plantas cultivadas em vasos, sob solo alagado e bem drenado durante 14 dias. O alagamento reduziu a área foliar específica e a alocação de biomassa para as raízes em todas as três espécies e aumentou a senescência foliar em B. brizantha e B. decumbens. O alagamento reduziu a taxa de crescimento relativo em B. brizantha e B. decumbens, mas não em B. humidicola. A taxa de elongação foliar não foi afetada pelo alagamento em B. decumbens e B. humidicola, mas diminuiu em B. brizantha desde o primeiro dia de alagamento. A fotossíntese líquida e o conteúdo de clorofila foliar foram reduzidos pelo alagamento em B. brizantha; no entanto, nenhum efeito do alagamento pôde ser detectado nas outras espécies. Em todas as espécies, existiu uma estreita relação entre as taxas de fotossíntese líquida e a condutância estomatal. Esses resultados mostram que as espécies estudadas diferem quanto à tolerância ao alagamento. B. brizantha é intolerante, B. decumbens é moderadamente tolerante e B. humidicola é tolerante. Em virtude de a taxa de elongação foliar ter sido imediatamente afetada somente em B. brizantha, este parâmetro pode ser empregado como um mecanismo de detecção prematura da tolerância ao alagamento em Brachiaria spp.Termos para indexação: Brachiaria brizantha, Brachiaria decumbens, Brachiaria humidicola, área foliar, clorofila, fotossíntese, biomassa, taxa de crescimento.

Nutrient enrichment is increasingly affecting many tropical ecosystems, but there is no information on how this affects tree biodiversity. To examine dynamics in vegetation structure and tree species biomass and diversity, we annually remeasured tree species before and for six years after repeated additions of nitrogen (N) and phosphorus (P) in permanent plots of abandoned pasture in Amazonia. Nitrogen and, to a lesser extent, phosphorus addition shifted growth among woody species. Nitrogen stimulated growth of two common pioneer tree species and one common tree species adaptable to both high- and low-light environments, while P stimulated growth only of the dominant pioneer tree Rollinia exsucca (Annonaceae). Overall, N or P addition reduced tree assemblage evenness and delayed tree species accrual over time, likely due to competitive monopolization of other resources by the few tree species responding to nutrient enrichment with enhanced establishment and/or growth rates. Absolute tree growth rates were elevated for two years after nutrient addition. However, nutrient-induced shifts in relative tree species growth and reduced assemblage evenness persisted for more than three years after nutrient addition, favoring two nutrient-responsive pioneers and one early-secondary tree species. Surprisingly, N + P effects on tree biomass and species diversity were consistently weaker than N-only and P-only effects, because grass biomass increased dramatically in response to N + P addition. The resulting intensified competition probably prevented an expected positive N + P synergy in the tree assemblage. Thus, N or P enrichment may favor unknown tree functional response types, reduce the diversity of coexisting species, and delay species accrual during structurally and functionally complex tropical rainforest secondary succession.