Nutrient Acquisition Strategies Augment Growth in Tropical N2‐Fixing Trees in Nutrient‐Poor Soil and Under Elevated CO2

Tropical forests play a dominant role in the global carbon (C) cycle, and models predict increases in tropical net primary productivity (NPP) and C storage in response to rising atmospheric carbon dioxide (CO2) concentrations. The extent to which increasing CO2 will enhance NPP depends in part on the availability of nitrogen (N) and phosphorus (P) to support growth. Some tropical trees can potentially overcome nutrient limitation by acquiring N via symbiotic dinitrogen (N2) fixation, which may provide a benefit in acquiring P via investment in N‐rich phosphatase enzymes or arbuscular mycorrhizal (AM) fungi. We conducted a seedling experiment to investigate the effects of elevated CO2 and soil nutrient availability on the growth of two N2‐fixing and two non‐N2‐fixing tropical tree species. We hypothesized that under elevated CO2 and at low nutrient availability (i.e., low N and P), N2 fixers would have higher growth rates than non‐N2 fixers because N2 fixers have a greater capacity to acquire both N and P. We also hypothesized that differences in growth rates between N2 fixers and non‐N2 fixers would decline as nutrient availability increases because N2 fixers no longer have an advantage in nutrient acquisition. We found that the N2 fixers had higher growth rates than the non‐N2 fixers under elevated CO2 and at low nutrient availability, and that the difference in growth rates between the N2 and non‐N2 fixers declined as nutrient availability increased, irrespective of CO2. Overall, N2 fixation, root phosphatase activity, and AM colonization decreased with increasing nutrient availability, and increased under elevated CO2 at low nutrient availability. Further, AM colonization was positively related to the growth of the non‐N2 fixers, whereas both N2 fixation and root phosphatase activity were positively related to the growth of the N2 fixers. Though our results indicate all four tree species have the capacity to up‐ or down‐regulate nutrient acquisition to meet their stoichiometric demands, the greater capacity for the N2 fixers to acquire both N and P may enable them to overcome nutritional constraints to NPP under elevated CO2, with implications for the response of tropical forests to future environmental change

Remotely Sensed Canopy Nitrogen Correlates With Nitrous Oxide Emissions in a Lowland Tropical Rainforest

Tropical forests exhibit significant heterogeneity in plant functional and chemical traits that may contribute to spatial patterns of key soil biogeochemical processes, such as carbon storage and greenhouse gas emissions. Although tropical forests are the largest ecosystem source of nitrous oxide (N2O), drivers of spatial patterns within forests are poorly resolved. Here, we show that local variation in canopy foliar N, mapped by remote‐sensing image spectroscopy, correlates with patterns of soil N2O emission from a lowland tropical rainforest. We identified ten 0.25 ha plots (assemblages of 40–70 individual trees) in which average remotely‐sensed canopy N fell above or below the regional mean. The plots were located on a single minimally‐dissected terrace (km2) where soil type, vegetation structure and climatic conditions were relatively constant. We measured N2O fluxes monthly for 1 yr and found that high canopy N species assemblages had on average three‐fold higher total mean N2O fluxes than nearby lower canopy N areas. These differences are consistent with strong differences in litter stoichiometry, nitrification rates and soil nitrate concentrations. Canopy N status was also associated with microbial community characteristics: lower canopy N plots had two‐fold greater soil fungal to bacterial ratios and a significantly lower abundance of ammonia‐oxidizing archaea, although genes associated with denitrification (nirS, nirK, nosZ) showed no relationship with N2O flux. Overall, landscape emissions from this ecosystem are at the lowest end of the spectrum reported for tropical forests, consist with multiple metrics indicating that these highly productive forests retain N tightly and have low plant‐available losses. These data point to connections between canopy and soil processes that have largely been overlooked as a driver of denitrification. Defining relationships between remotely‐sensed plant traits and soil processes offers the chance to map these processes at large scales, potentially increasing our ability to predict N2O emissions in heterogeneous landscapes

Sullivan2015GHG_rawdataforpublication

Soil greenhouse gas fluxes (CO2, CH4, N2O) and other environmental characteristics used in our analysis: Soil temperature, water potential, gravimetric water content, soil ammonium and nitrate concentrations, and soil available phosphorus concentrations

Data from: Proximate controls on semiarid soil greenhouse gas fluxes across 3 million years of soil development

Soils are important sources and sinks of three greenhouse gases (GHGs): carbon dioxide (CO2), methane (CH4), and nitrous oxide (N2O). However, it is unknown whether semiarid landscapes are important contributors to global fluxes of these gases, partly because our mechanistic understanding of soil GHG fluxes is largely derived from more humid ecosystems. We designed this study with the objective of identifying the important soil physical and biogeochemical controls on soil GHG fluxes in semiarid soils by observing seasonal changes in soil GHG fluxes across a three million year substrate age gradient in northern Arizona. We also manipulated soil nitrogen (N) and phosphorus availability with 7 years of fertilization and used regression tree analysis to identify drivers of unfertilized and fertilized soil GHG fluxes. Similar to humid ecosystems, soil N2O flux was correlated with changes in N and water availability and soil CO2 efflux was correlated with changes in water availability and temperature. Soil CH4 uptake was greatest in relatively colder and wetter soils. While fertilization had few direct effects on soil CH4 flux, soil nitrate was an important predictor of soil CH4 uptake in unfertilized soils and soil ammonium was an important predictor of soil CH4 uptake in fertilized soil. Like in humid ecosystems, N gas loss via nitrification or denitrification appears to increase with increases in N and water availability during ecosystem development. Our results suggest that, with some exceptions, the drivers of soil GHG fluxes in semiarid ecosystems are often similar to those observed in more humid ecosystems

Biogeochemical Recuperation of Lowland Tropical Forest During Succession

High rates of land conversion and land use change have vastly increased the proportion of secondary forest in the lowland tropics relative to mature forest. As secondary forests recover following abandonment, nitrogen (N) and phosphorus (P) must be present in sufficient quantities to sustain high rates of net primary production and to replenish the nutrients lost during land use prior to secondary forest establishment. Biogeochemical theory and results from individual studies suggest that N can recuperate during secondary forest recovery, especially relative to P. Here, we synthesized 23 metrics of N and P in soil and plants from 45 secondary forest chronosequences located in the wet tropics to empirically explore (1) whether there is a consistent change in nutrients and nutrient cycling processes during secondary succession in the biome; (2) which metrics of N and P in soil and plants recuperate most consistently; (3) if the recuperation of nutrients during succession approaches similar nutrient concentrations and fluxes as those in mature forest in ~100 yr following the initiation of succession; and (4) whether site characteristics, including disturbance history, climate, and soil order are significantly related to nutrient recuperation. During secondary forest succession, nine metrics of N and/or P cycling changed consistently and substantially. In most sites, N concentrations and fluxes in both plants and soil increased during secondary succession, and total P concentrations increased in surface soil. Changes in nutrient concentrations and nutrient cycling processes during secondary succession were similar whether mature forest was included or excluded from the analysis, indicating that nutrient recuperation in secondary forest leads to biogeochemical conditions that are similar to those of mature forest. Further, of the N and P metrics that recuperated, only soil total P and foliar δ15N were strongly influenced by site characteristics like climate, soils, or disturbance history. Predictable nutrient recuperation across a diverse and productive ecosystem may support future forest growth and could provide a means to quantify successful restoration of ecosystem function in secondary tropical forest beyond biomass or species composition

Proximate controls on semiarid soil greenhouse gas fluxes across 3 million years of soil development

© 2015 Springer International Publishing Switzerland Soils are important sources and sinks of three greenhouse gases (GHGs): carbon dioxide (CO2), methane (CH4), and nitrous oxide (N2O). However, it is unknown whether semiarid landscapes are important contributors to global fluxes of these gases, partly because our mechanistic understanding of soil GHG fluxes is largely derived from more humid ecosystems. We designed this study with the objective of identifying the important soil physical and biogeochemical controls on soil GHG fluxes in semiarid soils by observing seasonal changes in soil GHG fluxes across a three million year substrate age gradient in northern Arizona. We also manipulated soil nitrogen (N) and phosphorus availability with 7 years of fertilization and used regression tree analysis to identify drivers of unfertilized and fertilized soil GHG fluxes. Similar to humid ecosystems, soil N2O flux was correlated with changes in N and water availability and soil CO2 efflux was correlated with changes in water availability and temperature. Soil CH4 uptake was greatest in relatively colder and wetter soils. While fertilization had few direct effects on soil CH4 flux, soil nitrate was an important predictor of soil CH4 uptake in unfertilized soils and soil ammonium was an important predictor of soil CH4 uptake in fertilized soil. Like in humid ecosystems, N gas loss via nitrification or denitrification appears to increase with increases in N and water availability during ecosystem development. Our results suggest that, with some exceptions, the drivers of soil GHG fluxes in semiarid ecosystems are often similar to those observed in more humid ecosystems