Nitrogen deposition in tropical forests from deforestation and savanna fires

Running title: Fire effects on tropical N fluxes

Yang Chen, Department of Earth System Science, University of California, Irvine, CA 92697, USA James T. Randerson, Department of Earth System Science, University of California, Irvine, CA 92697, USA Guido R. van der Werf, Faculty of Earth and Life Sciences, VU University, 1081HV Amsterdam, Netherlands Douglas C. Morton, NASA Goddard Space Flight Center, Biospheric Sciences Branch, Code 614.4, Greenbelt, Maryland USA 20771 Prasad S. Kasibhatla, Nicholas School of the Environment, Duke University, Durham, NC 27708, USA

Kewords: atmospheric transport, pyro-denitrification, nitrogen limitation, Hadley circulation, tropical biomes, the global carbon cycle, and biomass burning Category of article: Primary Research Article

Corresponding author Yang Chen Email: yang.chen@uci.edu Phone: 1-949-824-0597 Department of Earth System Science, University of California, Irvine, CA 92697

Manuscript prepared for Global Change Biology June 11, 2009

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Abstract
Tropical forests account for nearly half of global net primary production (NPP) and may contribute substantially to contemporary and future land carbon (C) sinks. We used satellite-derived estimates of global fire emissions and a chemical transport model to estimate atmospheric nitrogen (N) fluxes from deforestation and savanna fires in tropical ecosystems. N emissions and deposition led to a substantial net transport of N equatorward, from savannas and areas undergoing deforestation to tropical forests. On average, N emissions from fires were equivalent to approximately 28% of biological N fixation (BNF) in savannas (4.8 kg N ha-1 yr-1) and 38% of BNF from ecosystems at the deforestation frontier (9.1 kg N ha-1 yr-1). N deposition occurred in interior tropical forests at a rate equivalent to 4% of their BNF (1.1 kg N ha-1 yr-1). This percentage was highest for African tropical forests in the Congo Basin (16%; 3.7 kg N ha-1 yr-1) owing to equatorward transport from northern and southern savannas. These results suggest that land use change, including deforestation fires, may be enhancing nutrient availability and carbon sequestration in nearby tropical forest ecosystems.

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INTRODUCTION
Nitrogen (N) is an essential element in Earth's atmosphere, biosphere, and hydrosphere (Galloway et al., 2003). The majority of N in the atmosphere is N2, which is biologically unavailable to most organisms. N2 must be converted to reactive N (Nr), which includes inorganic oxidized forms (e.g., NO, NO2, and HNO3), inorganic reduced forms (NH3 and NH4+), and organic forms (e.g., amino acids and urea), prior to its use by microbes and plants. N2 to Nr conversion (N fixation) occurs during electrical discharges in lightning and by some bacteria and algae via a process known as biological nitrogen fixation (BNF) (Cleveland et al., 1999). Humans have more than doubled the flux of N2 to Nr through fossil fuel use, industrial N fertilizer production, and cultivation of N fixing leguminous crops (Vitousek et al., 1997; Smil, 1999; Galloway et al., 2004, 2008). Atmospheric transport and deposition of N emissions substantially changes N availability in terrestrial ecosystems. Major effects of N deposition have been documented in terrestrial and marine ecosystems (e.g., Fenn et al., 2003). Since most terrestrial ecosystems are N-limited, N addition to these ecosystems stimulates net primary production (NPP) (LeBauer & Treseder, 2008) and may enhance carbon (C) storage by this mechanism (e.g., Holland et al., 1997, Reay et al., 2008). However, chronic N enrichment may lead to loss of C by increasing decomposition in organic soils (Mack et al., 2004) or in soil carbon components that have decadal turnover times (Neff et al., 2002). In some ecosystems, N addition also reduces microbial biomass and decreases soil respiration (Treseder, 2008). N deposition can also influence ecosystem function by means of acidification, eutrophication (Rabalais, 2002), and loss of species diversity (Wilson and Tilman, 2002; Suding et al., 2005; Phoenix et al.,

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2006). Much of our knowledge of the N cycle and the consequences of N enrichment are from temperate ecosystems, where industrial and agricultural modifications of N availability are substantial. Tropical ecosystems respond in different ways to perturbations in N cycling (Matson et al., 1999; Davidson et al., 2004) and important uncertainties remain with respect to our understanding of N deposition, fixation, and loss pathways in these ecosystems. With implementation of stricter air quality standards, N emission and deposition are likely to slow down in temperate regions, particularly in developed countries (Lamarque et al., 2005; Dentener et al., 2006). In contrast, N emission and deposition in tropical and subtropical regions should increase (Galloway et al., 1994). Since tropical ecosystems account nearly half of global NPP (Field et al., 1998) and may contribute substantially to contemporary and future land C sinks (Friedlingstein et al., 2006; Stephens et al., 2007), it is imperative to identify and quantify the consequences of changing levels of N deposition and N cycling in tropical ecosystems. Fires in savanna and at the deforestation frontier are a major source of air pollution in the tropics (Crutzen and Andreae, 1990). Nitrogen oxide (NO) and ammonia (NH3) are the primary Nr gases emitted from these fires. They are converted to other Nr gases and to particulate species (NO3-, NH4+, and organic aerosols) by means of multiple reaction pathways (Crutzen and Andreae, 1990; Atkinson, 2000). The ultimate fate of Nr in the atmosphere is removal by wet and dry deposition. Gas and aerosol components of dry deposition are important in many regions (Dentener et al., 2006). Regional variations in emissions and atmospheric transport of Nr have the potential to create strong gradients of

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2001).. 2001.. Deutsch et al. which combines satellite observations of burned area (Giglio et al. We performed 10 years (1997–2006) of global-scale simulations but focus our analysis and discussion on tropical ecosystems.. 1990. These fluxes were then compared with terrestrial and marine BNF fluxes (Cleveland et al. In this study. van der Werf et al. 1991). as some is retained within the ecosystem in combusted residues (approximately 20%) and the remainder (another 30–40%) is transformed directly into N2 during the combustion processes (Lobert et al. We estimated N2 emissions from biomass burning using data derived from a chamber burning experiment (Kuhlbusch et al.N deposition across both terrestrial and ocean ecosystems. We explored the loss of N by emissions from biomass burning and the gain of Nr by deposition from the atmosphere. 1991)... Lee et al.. 2002). We validated some of our modeled fluxes by comparing our results with published observations of N deposition. 1999. We also estimated the net N flux from fire emissions in different tropical ecosystems... The N2 emission represents a net loss of N from the biosphere– contrasting with Nr fluxes that instead lead to redistribution of Nr across different ecosystems. Kuhlbusch et al. we assessed the contribution of fires to N cycling in tropical ecosystems. We estimated fire emissions using the Global Fire Emission Database version 2 (GFEDv2. 5 . The transport and deposition of N were simulated by using the GEOS-Chem global chemical transport model (CTM) (Bey et al. 2006) and a biogeochemical model constrained by other satellite data to estimate fuel loads. Emissions of NO and NH3 probably account for only 40-50% of the N present in fuels prior to combustion. 2006)..

fuel loads (FL).METHODS Nr emissions from biomass burning The GFEDv2 product consists of 1°×1° gridded monthly burned area (BA). Burned area was derived using Moderate Resolution Imaging Spectroradiometer (MODIS) active fire and burned area datasets as described by Giglio et al. required a small negative adjustment (from 0-22% depending on atmospheric inversion approach) to match Measurements of Pollution in the Troposphere (MOPITT) observations during 2000-2006 (van der Werf et al. combustion completeness (CC). 2006). Comparison with atmospheric carbon monoxide (CO) observations provides some validation of carbon emissions. and emissions factors (EMFs) for different gas and aerosol species (van der Werf et al.. 2006). Emission rates (E) for each gaseous N species were calculated by applying EMFs to fire-emitted carbon fluxes from CASA: E = EMF × BA × FL × CC (1) Organic dry matter (DM) in the fuel was assumed to be comprised of 45% carbon.. for example. CO anomalies from GFEDv2 in the GEOS-CHEM model had a mean correlation of 0. Emission factors. defined as the emission of species (g) per 1 kg burned DM. (2006). Fuel loads were estimated using the Carnegie-Ames-Stanford-Approach (CASA) biogeochemical model constrained by additional satellite observations of fractional tree cover and the fraction of absorbed photosynthetically active radiation by plant canopies (van der Werf et al.79 with observations from NOAA Global Monitoring Division (GMD) stations in the Northern Hemisphere 6 . 2008). GFEDv2 CO emissions in equatorial Asia. were obtained from Andreae and Merlet (2001).. 2003.

they are of sufficient quality to study the impact of fire-emitted species on atmospheric composition.59 with GMD stations in the Southern Hemisphere (Kai et al. Nr can be emitted into the atmosphere in different chemical forms and physical phases. for northern and southern hemispheres. Kuhlbusch et al. Similarly. and acetonitrile) is small compared with uncertainties associated with our estimates of burned area (e.g.. NOx and NH3 account for the majority (greater than 90%) of Nr emissions from biomass burning (Andreae and Merlet. van der Werf et al.86 and 1. CO/CO2 emission ratio 7 . By burning different biomass materials in a stainless steel chamber. the ratio of modeled to observed standard deviations of CO anomalies from GMD was 0. 2008).g. 1990). The low bias we expect from neglecting the remaining Nr emissions (e. (1991) showed that the missing N is mostly molecular N (N2).and 0. These results indicate that while emissions estimates are still uncertain.. Giglio et al. This experiment also showed that N2 is formed primarily during the flaming stage and thus the total amount of N2 production (as a fraction of fuel N) depends on fire intensity. Nr to N2 conversion during biomass burning A large gap has been observed in the N balance between its content in the fuel and the sum of measured N in gaseous emissions and what remains after combustion in the ash (Lobert et al. N2O... we used the sum of gaseous NOx and NH3 emissions to represent total Nr emissions from biomass burning. which accounts for approximately one third of fuel N. 2009). In our model simulations and calculations..g..09. 2006) and total carbon emissions (e. 2001). respectively.. HCN.

Analysis of the Kuhlbusch et al. transformation. (1991) measurements showed that the N2-N to Nr-N ratio of fire emissions was inversely related to its CO/CO2 ratio (see Fig. biofuel.5°(longitude) vegetation map of these three types (Andreae and Merlet. 1998)..5° horizontal resolution and 30 vertical layers between the surface and 0. and deposition of biomass burning emitted N species. For each grid cell. we estimated the CO/CO2 ratio for three different fire types: savanna and grassland fires. S1 in Supporting Information). and extratropical forest fires using the emission factors of CO and CO2 listed in Table 1 of Andreae and Merlet (2001). In this study. the GFEDv2 Nr emission rate was scaled by this ratio to approximate the fire emission rate of N2.. tropical forest fires. The N2-N to Nr-N ratio derived from this relationship was used to estimate global N2 emissions from biomass burning.01 hPa. A 2°(latitude) × 2. 2001) driven by assimilated meteorological observations from the Goddard Earth Observing System (GEOS) of the NASA Global Modeling and Assimilation Office (GMAO). The GEOS-Chem model includes a detailed description of tropospheric O3-NOx-hydrocarbon chemistry (Wang et al. aircraft. 2001) was used to derive the global gridded N2-N to Nr-N ratio. In this study.serves as a powerful indicator for the relative extent of flaming combustion and smoldering combustion. GEOS-Chem is a global 3-D CTM (Bey et al. Model simulations of N transport and deposition We used the GEOS-Chem model to calculate the transport. biomass burning. soil and fertilizer. we used version 08-01-01 of the model with 2°×2. and NH3 from biofuel. fossil fuel. fossil fuel. Emission sources of N include NOx from lightning. biomass burning. and 8 .

and N on sea salt).g. and another without these emissions. We performed GEOS-Chem full chemistry simulations over a 10-year period (1997-2006) using the GFEDv2 inventory. N2O5. peroxyacyl nitrate (PAN). vegetation. two sets of simulations were conducted: one with biomass burning emissions prescribed from GFEDv2. soils. and upper-bound) between BNF and evapotranspiration (ET) in terrestrial ecosystems. and wild animals). These tracers are transported by meteorology-driven advection and convection processes. After a three-month spin-up period with the same initial conditions. and NH3) or aerosol phase (NH4+. NOx. HNO3... central. (2004) argued that the true N fixation rate lies at the lower end of this 9 . (1999) reviewed published studies and generated three linear regressions (conservative. N on sulfate.g.5° model grid cell. N emissions from other sources were obtained from climatological inventories which represent contemporary (circa 2000) emissions. The difference between these two simulations represents the effect of biomass burning emissions on N transport and deposition. and finally removed by the action of precipitation (wet deposition) or by contact with surface or vegetation (dry deposition). which resolves the interannual variability of biomass burning emissions. We recorded monthly mean deposition rates for each N species through dry and wet deposition..natural sources (e. Biological N fixation Cleveland et al. Wet deposition included convective and large-scale components (Liu et al. Total deposition rate of N from biomass burning was then calculated for each 2°×2. Galloway et al. N tracers are present in gas phase (e. 2001).

we therefore used the mean of central and conservative regressions between BNF and ET from Cleveland et al. we assumed that the mean of these two rates represented the N fixation rate over other tropical oceans. 2002) and Pacific (Deutsch et al. 2001)... Estimates of global ocean BNF range from 60 to 200 Tg N yr-1 (Duce et al. In this study. we used reported estimates of BNF rates over tropical Atlantic (Lee et al. 2008). In this study. (1999) to estimate N fixation rates over land. The global ET map we used was developed using satellite data and a biometeorological approach by Fisher et al. due to the inherent biases noted in plot-scale studies. Large uncertainty remains in the estimation of regional marine N fixation (Mahaffey et al. Due to a dearth of data for other regions. (2008). 2005). 10 .range...

We compared these observations with modeled deposition fluxes. About 75% of the wet deposition rates and 80% of the dry deposition rates agreed within ±50% with the observations. The location and time period of these measurements. We only considered observations from 1996 to 2006 to ensure overlap with the period of our satellite-derived fire emissions time series.. 2005. are summarized in Table S1. Dentener et al. Here we compiled wet and dry N deposition measurements in tropical regions published in recent years. the model simulation corresponded reasonably well to wet and dry deposition observations at non-urban sites (Fig. Based on the original data. We differentiated urban and non-urban observations according to the description of the site in the original literature or the proximity of the measurement station to an urban center. which were sampled in grid cells corresponding to observations during the month when observations were available. The model somewhat underestimated NO3-N wet deposition and overestimated NH4-N dry deposition. fewer N deposition measurements exist for tropical land and ocean regions. we calculated wet and dry deposition rates for total oxidized N (NO3-N) and reduced N (NH4-N) with units of kg N ha-1 yr-1. as well observed and modeled deposition rates. 11 . For locations that had measurements only during 1996. Altogether. Overall. we compared the deposition rates with our 10-year (1997–2006) mean model results. This agreement was similar to previous comparisons of N deposition between models and observations (Lamarque et al. 2006).RESULTS Validation of wet and dry deposition Relative to Northern Hemisphere industrial regions. we reported wet and dry deposition rates from 26 non-urban stations. 1)..

Urban data were generally not representative of the 2°×2. Losses directly to N2 accounted for about half of the total N emissions flux. In contrast to non-urban sites. the model was generally biased low relative to the high urban data. Gross N emissions from fires Emissions of Nr and N2 by fires were 10. N deposition rates from biomass burning Fire emissions represented a considerable fraction of N deposition in the tropics (See Fig. 2a and Table 1).3 12 . eastern Bolivia.9 kg N ha-1 yr-1).4 Tg N yr-1 (6. so they were not included in our linear regressions. This fraction was slightly higher for savannas than for tropical forests because of more complete combustion (and lower CO/CO2 ratios in savannas). High emission rates occurred in African savannas. 1. S2) in South America and Southeast Asia. with these two biomes accounting for 67% of global fire N emissions (Fig.5° grid cells in the model. Deposition rates were highest near source regions .8 kg N ha-1 yr-1) in tropical forests. in southern Borneo in equatorial Asia (up to 9. and Paraguay in South America (up to 5. We note most of these urban measurement sites were located in Southeast Asia. as indicated by the r values shown in Fig.3 Tg N yr-1 (3. as well as in deforestation regions (See Fig.5 kg N ha-1 yr-1) in savannas and 5. and across the Brazilian state of Mato Grosso.Simulations were more strongly correlated with measurements for dry deposition than for wet deposition.with maxima on the perimeter of the Congo basin in Africa (up to 9. S3).7 kg N ha-1 yr-1). which were likely affected by nearby fossil fuel emissions (Fig. 1).

1998).. Separation of N deposition by its oxidation form is important for assessing N deposition effects on ecosystem function. parks and other protected areas) experience relatively high N deposition from nearby deforestation and savanna fires. 2008) and peatlands (Paulissen et al. 2c.g. 2b).kg N ha-1 yr-1) (Fig.. for example. probably as a consequence of increasing precipitation levels. These results imply that undisturbed forest and savanna ecosystems in these regions (e. although regionally the relative importance of these two pathways varied. Changes in N surface budgets caused by fire emissions The emission and deposition of N from fires led to a net loss of N from the terrestrial biosphere and a redistribution of N among ecosystems. Savanna ecosystems in Africa and South America and tropical deforestation frontiers in all three regions experienced a net loss of N (Figs. About 2/3 of the total N wet deposition occurred in the gas phase (NH3 and HNO3). Reduced N. Net gains at a 13 . Seasonal variations in N deposition were closely linked with the timing of fire emissions (Fig. causing changes in species composition in ecosystems such as heathlands (van der Berg et al. 3). can reduce base cation uptake by plants (de Graaf et al. 3).. 2004). Wet deposition accounted for a larger fraction of total deposition in Southeast Asia (57%) and Northern Hemisphere (NH) Africa (55%) than in South America (46%) or Southern Hemisphere (SH) Africa (39%). Total wet deposition was comparable to total dry deposition across the tropics. The relative importance of wet deposition as an atmospheric loss pathway increased towards the end of the fire season on all three continents (Fig. Model estimates of the total deposition rate for oxidized N and reduced N were similar. S2).

Most of the remainder was deposited in downwind tropical forests (e.7 kg N ha-1 yr-1 in savanna ecosystems (Table S2). For interior forests in Africa.g. with 55% of Nr emissions remaining within these ecosystems via deposition in nearby areas. corresponding to approximately 16% of BNF. Of the directly emitted Nr from African savanna fires. tropical forest fires had a net N loss equal to approximately 10% of BNF (2. Van der Werf et al. For savannas across all three continents. The net loss of N (Nr and N2) from savanna ecosystems was largest in Africa (Table 1). where fire return times were lower than on other continents (Giglio et al.. In interior tropical forests that were removed from the deforestation frontier (areas with fire emissions below our 500 kg C ha-1 yr-1 threshold)...19 Tg N yr-1 or 6. in the Congo basin) and over the tropical Atlantic Ocean (Fig.8 kg N ha-1 yr-1). Net fire losses were equal to approximately 39% of BNF in savanna ecosystems in Africa. Overall. In Southeast Asia.0 kg N ha-1 yr-1). Tropical forests received large N deposition inputs from savanna fires but also lost N through deforestation fires. the net atmospheric flux from fires was 0. Expressed as a percentage of BNF.7 kg N ha-1 yr-1). 2008). because a larger proportion of fire emissions were associated with deforestation in tropical forests. forests at the deforestation frontier in South America and Southeast Asia had a large net N loss at a regional scale from fire emissions. net losses were higher (35%). corresponding to 6. the net effect of remote tropical fires was an increase atmospheric deposition.regional scale occurred in interior tropical forests and over tropical oceans. 2006.46 Tg N yr-1 (3. 2b). net N losses at the deforestation frontier (38%) were 14 . only 53% was returned to savannas via deposition. for example. In contrast.83 Tg N yr-1 or 2. net fire N losses were equal to 28% of BNF (4.

even higher than in savanna ecosystems. received high N deposition fluxes – up to 3. Approximately 37% (2. owing to high levels of fire emissions in Africa and the prevalent westerly wind during the fire season (see Fig. 2 and Table 1). Evergreen broadleaf forests were distributed mostly between 5°N and 5°S and were bordered to the north and south by savannas (Fig. A portion of Nr emitted from savanna fires was deposited back in savannas.85 Tg N yr-1) of fire-emitted N was deposited over oceans. However.7 kg N ha-1 yr-1) from fires in Indonesia and Peninsular Malaysia. especially that for dry deposition. A small amount of the N from African fire emissions was transported across the Atlantic and deposited over South America. in particular. Redistribution of N between ecosystems in Africa Fires caused a net transport of N from savannas to interior forests in tropical Africa (Fig. 4a). Tropical oceans were a large recipient of N from fire emissions. 2009). a substantial amount of fire-emitted Nr was transported equatorward and 15 . 4b). The southern part of South China Sea also received substantial N deposition (up to 2.8 kg N ha-1 yr-1 (Fig. This pattern is consistent with analysis of lidar measurements from Manaus that show long range transport of smoke and dust from Africa (Ansmann et al. was similar to that of fire emissions but with a small equatorward shift in peak deposition in the NH. The N deposition pattern. The highest levels of Nr and N2 fire emissions occurred at ~7°N in the northern hemisphere and ~10°S in the southern hemisphere.. S4). The tropical Atlantic. 2). BNF was highest near the equator and decreased to the north and south with decreasing ET (Fig.

deposited in tropical forests. and deposition resulted in a net transport of N from savannas to forests in Africa (Fig. 4c). Prevailing winds in South America and Southeast Asia also caused transport of fire emissions toward interior (and relatively undisturbed) areas of the Amazon and the Indonesian archipelago (Fig. In NH winter. respectively. 2c). 4f and Fig. Convective lifting of air masses near the equator increased precipitation and. subsequently. This equatorward transport pattern was not unique to Africa. S4). The transport was reversed in NH summer. received relatively small N deposition fluxes from fire emissions. which corresponded to the NH fire season. prevailing winds were to the south (Fig. when winds to the north carried emissions from SH savannas to interior tropical forests. The Sahara and Kalahari deserts. The combined effects of biomass burning emissions. 16 . atmospheric transport. wet deposition. which were located upwind of the African savanna regions during the fire seasons in the northern and southern hemispheres.

Our results indicated a redistribution of N between different types of tropical ecosystems. 2009) and African (Lewis et al. Interior tropical forests gained N equivalent to 4% of their BNF. Lewis et al. Ciais et al... The Hadley circulation carried emissions from these fires equatorward. This atmospheric transport of N was closely related to the seasonality of fire emissions and meteorological patterns in tropics. A significant amount of N from this zonal transport was removed by wet and dry deposition over relatively undisturbed tropical forests.. fire N losses were equivalent to 28% of BNF in savannas and 38% of BNF for ecosystems at the deforestation frontier. S4). On average. 2004) of long-term increases in carbon storage observed within Amazonian (Phillips et al. 2009) forests. toward the Intertropical Convergence Zone (ITCZ) (See Fig. intact tropical forests. the prevailing winds carried fire emissions from savannas to tropical forest during both NH and SH fire seasons. The cross biome transport of N described here strengthens the case for increases in nutrient deposition serving as a key driver (e. 2006. Specifically.DISCUSSION AND CONCLUSIONS We estimated emissions and deposition of N from tropical fires. Although modeling studies suggest tropical forests have great potential to increase net primary production in response to elevated levels of carbon dioxide (Friedlingstein et al. Most fires occurred in savannas and in areas that were undergoing active deforestation. In tropical Africa. we found an equatorward transport of N from fires in savannas and at active deforestation frontiers to interior. a key uncertainty is whether nutrient availability can keep pace with photosynthesis to allow for long-term increases in 17 ...g.. the net transport of N from savannas to forests in Africa was larger than that observed in South America or Asia. 2008). As a consequence of widespread savanna burning in Africa.

management of deforested areas. Indeed. Luo et al. The impacts of these deposition fluxes for carbon accumulation may be largest in younger secondary forests where nitrogen and phosphorous limitation are most pronounced (Davidson et al.. In this context. the net effect of the advancing deforestation frontier is therefore to expand the area under fire management and to increase the proximity of savanna fires to highly productive interior forests. In parallel to the patterns shown here for N. Hungate et al. and land use in neighboring savanna regions may be enhancing nutrient availability in nearby tropical forest ecosystems– and thus increasing the sensitivity of these ecosystems to changing levels of atmospheric CO2. fire-emitted black carbon aerosols have been shown to contribute in important ways to phosphorous deposition in tropical forests within the Amazon (Mahowald et al. Our analysis shows how concentrated fire use for deforestation.. 2005).biomass (e. Other land use-related changes in nutrient cycling. free air carbon dioxide enrichment studies from mid-latitude forests show that rates of carbon accumulation in response to elevated levels of carbon dioxide increase when nutrient availability is enhanced (Oren et al.. 2004. Satellite-based fire detections suggest that fire use for pasture and cropland management sustains high fire activity following deforestation (Schroeder et al. In terms of assessing the net impact of land use change on global levels of atmospheric CO2. 2005). 2003. 2007).. may additionally increase the nutrient loading to forests near the deforestation frontier. increased nutrient loading by deposition of fire emissions in intact and secondary tropical forest ecosystems may partially offset some of the carbon losses associated with tropical deforestation. it is 18 . including the use of N fertilizer in agriculture after forest clearing and increases in dust. Davidson et al....g. 2001).. 2004).

In equatorial Asia after the 1997/1998 El Nino. However.important to further refine our understanding of the fire-emitted component of deforestation carbon losses (e. for example. particularly in coastal systems (Rabalais. for redistribution of nutrients.. ozone. 2007. these loss pathways are fundamentally different. changes in aerosol optical depth and diffuse light (Oliveira et al. increases in atmospheric N deposition by anthropogenic activity may be partially offset by lower rates of N fixation (Krishnamurthy et al. methane. 2009.. Mercado et al. including phytoplankton primary production and the health of coral reefs. 2005). 19 . 2002). 2007). Detailed analysis of the effect of atmospheric N deposition on marine ecosystems is complex for several reasons. Nevertheless. coral reef death was attributed to extraordinary red tide caused by deposition of iron from fire emissions (Abram et al.. Second. DeFries et al. First. 2009). currently there are not enough observations to derive a global map of ocean N fixation (Capone et al.. this deposition is relatively small (2%) compared to BNF and also with respect to N transport to surface waters by upwelling (Capone et al... Further work is needed to identify how nutrient loading from fires affects marine ecosystem function in these highly impacted areas.. Doughty et al.. 2005). 2008). N is generally considered a primary limiting nutrient for phytoplankton biomass accumulation in marine ecosystems. Averaged over the whole tropical ocean. In terms of atmospheric CO2 levels. A large portion (~37%) of N from savanna and forest fires is carried to and deposited over tropical oceans (Table 1).g. However. decomposition and fire carbon loss pathways from deforestation may not lead to substantially different outcomes. 2003). and human health impacts.. the ratio is much higher in coastal regions in Africa and Southeast Asia (Table 1).

More detailed measurements of N losses to N2 during fires also are needed to reduce uncertainties associated with pyrodenitrification. Although considerable effort has gone into improving our understanding of the deposition of chemical species in tropics (e. the Deposition of Biogeochemically Important Trace Species (DEBITS) network.gov/DEBITS. 20 .noaa. http://www. Finally. an improved understanding of the fate of N lost during deforestation and N cycling during post-deforestation land use may enable us to better quantify human impacts on N cycling in tropical ecosystems..Here we show that N emissions from savanna and deforestation fires lead to deposition in tropical forest ecosystems as a consequence of atmospheric transport associated with the ITCZ.igac.php).g. Better measurements are required to improve the model simulation of N transport and deposition. measurements of tropical N deposition are still sparse compared to the density of available observations in temperate regions. including N deposition measurements near deforestation regions and improved estimates of fire emissions associated with deforestation. A quantitative understanding of this transport pathway is currently limited by several different types of observations.

Kasibhatla P. was partially supported by the grant NNX08AL03G from Duke University.S.ACKNOWLEDGEMENTS This work was funded by NASA NNX08AF64G. 21 . The GEOS-Chem model is managed by the Atmospheric Chemistry Modeling group at Harvard University with support from the NASA Atmospheric Chemistry Modeling and Analysis Program.

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49 0.01 10.29 15.28 0.31 1.04 0.83 20.15 1.63 0 1. (1999).46 13.03 0 Fire emissions of N2 Tg N yr-1 0.07 0.21 -0.51 0.87 -2.04 0.87 2. In this study.70 0.75 2.16 0.21 0.44 27.70 2.13 0..82 -0.64 0.51 8.80 28.31 -0.61 0.80 0.69 9.21 0. Area Continent Region Mha Tropical forest Interior Africa 3 Biological Nitrogen Fixation2 Tg N yr-1 5.34 0 0.32 0 2.22 0.55 3.59 0.19 1.19 0. ‘Total tropics’ refers to regions within 30°S-30°N.22 4.36 1.28 -1.39 0. 2 3 Estimated using the mean of central and conservative regressions between BNF and ET as given in Cleveland et al.Table 1. Here it is defined as region with forest cover greater than 10% and the fraction of forest loss during 2000-2005 (Hansen et al.94 2.90 0.63 0.01 2.81 0.12 0.85 % of BNF -3 16 -5 -39 6 -4 1 -21 -10 3 -35 8 -60 -35 6 -10 4 -38 -28 2 288 123 5 921 2966 693 390 64 397 1494 304 16 56 112 2523 1409 529 145 1610 19530 Frontier4 Savanna Ocean Tropical forest Interior South America Frontier Savanna Ocean Tropical forest Interior Southeast Asia Frontier Savanna Ocean Tropical forest Interior Total tropics Frontier Savanna Ocean 1 The domains of tropical regions in Africa. South America.12 0. ‘Interior’ refers to tropical forest region that is rarely affected by human activities. The values are 10 year (1997-2006) averaged.56 -1. 31 . 2.83 0.54 0.11 Fire emissions of Nr Tg N yr-1 0.21 0.01 0.03 -0.52 0.32 -7.67 0.66 0.24 0.07 -0.52 12.32 0 2.62 0.17 0.86 5. N fluxes from fires over tropical regions1.46 -0. 2008) greater than 1%.69 0.54 0.67 0.11 15.88 5.01 4.5° grid cell with fraction of evergreen broad leaf forest (from MODIS 500 m land cover product) over 70% and the 10-year average fire emission rate (from GFEDv2) smaller than 500 kg C ha-1 yr-1.00 0 0.12 0.04 0.42 2.01 0.85 % of BNF 20 19 13 14 6 4 2 8 5 3 13 13 18 9 6 9 6 12 10 2 Net fire N flux Tg N yr-1 -0.01 -6. and Southeast Asia are shown in Fig. 4 ‘Frontier’ refers to tropical forest region undergoing substantial anthropogenic deforestation.70 0.58 0 1. it is defined as 2°×2.34 1.41 0 Deposition of fireemitted N Tg N yr-1 1.66 120.05 0.07 0.01 4.

Filled diamonds. (c) meridional wind speed (m/s) at 850hPa. (b) biological N fixation (kg N ha-1 yr-1) estimated from Cleveland et al. Red and blue solid lines show linear regression fits for NO3-N and NH4-N deposition rates. circles. The values are in kg N ha-1 yr-1. South America. and (f) net fire N and Nr fluxes (kg 32 . Blue and red colors represent N addition to and loss from terrestrial ecosystems. and triangles represent data for Africa. Gray points are observation stations near urban areas. Figure 3: Seasonal variability of total dry and wet deposition (Tg N yr-1) of N from biomass burning over NH Africa. Solid pink boxes define the domains of tropical Africa (AFR). with northward wind denoted as positive. 2). Dashed lines indicate 1:1. and 2:1 relationships. South America. Figure 2: Simulated 10-year average (a) N emissions from fires (the sum of Nr and N2). The net fire N flux includes both losses from Nr and N2 emissions and gains from N deposition. 1:2. The values are the 10 year (1997-2006) mean from the GEOS-Chem simulation. (2008). Figure 4: Latitudinal distributions of mean (a) fraction of forest and savanna cover. SH Africa. tropical South America (SAM). (1999) using a global map of ET from Fisher et al. and (c) net fire N flux with respect to the surface. and Southeast Asia (the domains are defined in Fig. Red and blue points represent deposition rates for oxidized N (NO3-N) and reduced N (NH4-N). (e) N deposition from biomass burning. respectively. and in Southeast Asia.FIGURE CAPTIONS Figure 1: Comparison of observed and modeled N wet and dry deposition rates (kg N ha-1 yr-1) in tropical regions. and tropical Southeast Asia (SEA) used in this paper. (b) N deposition from biomass burning. respectively. (d) N emissions from fires.

NH savanna. is averaged over February (representing fire season in NH) and August (representing fire season in NH). The values are 10 year (1997-2006) averaged. This domain covers. derived from GEOS-4 reanalysis data.N ha-1 yr-1) for tropical African region (30°S-30°N. 33 . tropical forest. and the Karahari desert in Southwest Africa. The meridional wind speed. The net fire Nr flux includes losses from Nr emissions and gains from N deposition. SH savanna. the Sahara desert. 10°E-30°E). from north to south.

Figure 1 34 .

Figure 2 35 .

Figure 3 36 .

Figure 4 37 .

SUPPORTING INFORMATION The following Supporting Information is available for this article: Figure S1 N2-N/Nr-N ratio vs. CO/CO2 ratio Figure S2 Forest cover loss due to deforestation and tropical ecosystem masks Figure S3 Biomass emission and wind vectors Table S1 Summary of N deposition observations Table S2 N fluxes from fires in tropical regions– expressed in units per unit of land area 38 .

dashed. 39 . The dash-dotted.. and extratropical forest fires. respectively. Data are based on results from a chamber experiment (Kuhlbusch et al. excluding two pine needles samples (gray points) because of the limited abundance of conifer plant functional types in tropical regions. We also excluded three wood points below the N2 detection limit. 1991). and dotted vertical lines represent the CO/CO2 values for savanna fires.Figure S1. N2-N/Nr-N ratio as a function of CO/CO2 ratio. tropical forest fires.

(c) Tropical savanna cover fraction (%). The tropical ecosystem cover fractions and masks were derived from MODIS 500 m land cover product.5º resolution.Figure S2. The original data were compiled by Hansen et al. (a) Deforestation within the humid tropics during 2000-2005.5º×0. (d) Tropical ecosystem masks in 2º×2. 40 . shown as the percent loss within each 0. (b) Tropical forest cover fraction (%). (2008).5º cell.

41 . The values are 10 year (1997-2006) averaged. Latitudinal distribution of Nr emission and deposition rates (kg N ha-1 yr-1) from fire and other sources. Inserted plots show the mean values and fire contribution fractions for three tropical regions as defined in Fig.Figure S3. 2.

Red shaded areas depict the Nr emissions (kg N ha-1 yr-1) from biomass burning for these months (10-year average). The GEOS-4 wind vectors at 850hPa in tropical regions for February and August. 42 . The full length of an arrow represents a wind speed of ~20m/s.Figure S4. 2004.

26 11.56 0. List of wet and dry N deposition observations and corresponding simulated values at non-urban sites in tropical regions..83 2.03 0.71 0.29 0.93 0... 1998-2003 Apr-Jul.29 2.00 1. Indonesia 0º12' S 100º19' E 2000 – 2005 4.10 0.88 0.06 2.60 1. 2008 Da Rocha et al. Niger Katibougou.08 1. 2009 Trebs et al..08 2. Indonesia Asia Kototabang.06 1.73 4. Brazil Andean montane forest..cc EANET 3º10' N 13º20' N 23º00' S 6º N 25º S 13º20' N 12º30' N 10º04' S 4º S 21º48' S 5º21' S 22º54' S 4º S 4º S 10º04' S 10º04' S 10º04' S 4º28' N 11º58' E 2º25' E 30º02' E 5º W 31º38' E 2º25' E 8º5' W 61º56' W 79º05' W 48º11' W 53º41' W 47º03' W 79º05' W 79º05' W 61º56' W 61º56' W 61º56' W 101º22' E 09/1996 – 04/2000 06/1996 – 10/1996 1986 . value in kg N ha -1 month-1 Trebs et al. 2006. Brazil Andean montane forest.13 0.23 0. value in kg N ha -1 month-1 Ayers et al.73 1.Table S1..53 1..83 1.88 1.90 0.2002 07/1999-06/2002 06/1994-09/2005 1997-2005 Sep-Nov.28 0. The shaded rows represent dry deposition measurements and other rows represent wet deposition measurements. 2009 Galy-Lacaux et al.07 3.97 (kg N ha-1 yr-1) 3.09 0.04 0. Ecuador Araraquara.00 0. 1998 Mphepya et al.84 0.62 2. Ecuador Andean montane forest.24 0. 2008 Boy et al.14 6º15' S 106º34' E 2001 .07 2.67 1.20 0. Brazil Rondônia.. Ecuador Rondônia. Updated in Galy-Lacaux et al.83 Sigha-nkamdjou et al..89 1..63 0.95 2.86 0.52 0. 2002 05/1998-04/2003 04/1999-02/2001 1999 08/1997-07/1998 Aug-Mar.44 2. 2003 Galy-Lacaux et al.99 1. NO3-N (obs) Region Location Latitude Longitude Time (kg N ha-1 yr-1) Zoétélé.67 0. Cameroon Banizoumbou.78 0. 1998-2003 09/2002 10/2002 11/2002 1996 1. 2005 Yoboué et al. 2006.1999 1995 . Mali Rondônia.. 2005 Lara et al.42 0..20 0..18 1.56 2. 2002 Acid deposition Monitoring Network in East Asia NO3-N (mod) NH4-N (obs) NH4-N (mod) Reference 43 . 2007.22 0. RSA Banizoumbou.31 0.60 0.. Côte d'Ivoire Skukuza.02 2.80 1.80 0. Malaysia Southeast Serpong. RSA Africa Lamto..02 (EANET).57 0.26 1. 2006 Boy et al.2005 8. French Guyana South America Piracicaba River Basin.25 0.05 2. Brazil Petit Sault. Brazil Rondônia. Averaged over 4 sites Boy et al. 2008 Trebs et al.92 2. 2006.eanet.83 1..57 4.60 0.84 1. Niger Louis Trichardt.28 0... 2006 Galy-Lacaux et al. 2004. Brazil Tanah Rata..10 0.38 (kg N ha-1 yr-1) 4.63 1.86 0.75 0. Data available online: http://www. value in kg N ha -1 month-1 Trebs et al.26 1. 2005 Mphepya et al.52 0.96 (kg N ha-1 yr-1) 1.97 2.09 0. 2009 Yoboué et al.58 1.81 0.

93 0. Malaysia Patumthani.53 2.23 1.31 1.58 4.59 2.16 1.Tanah Rata.74 10. Thailand Mae Hia.93 3. Thailand Vachiralongkorn Dam.48 3.55 1.69 EANET EANET EANET EANET EANET EANET 44 .31 2.71 1.95 6.62 0. Thailand Khao Lam.08 3.01 2.08 3. Thailand Hoa Binh.93 5.55 2.07 1. Vietnam 4º28' N 14º02' N 14º4' N 14º46' N 18º46' N 20º49' N 101º22' E 100º46' E 101º37' E 98º35' E 98º56' E 105º20' E 2000 – 2005 2000 – 2005 2000 – 2001 2002 – 2005 2001 – 2005 2000 .2005 2.66 1.01 5.

2 6.4 25.7 -1.5 1.8 1.4 25.1 -9. Area Continent Region Mha Tropical forest Interior Africa Frontier Savanna Ocean Tropical forest Interior South America Frontier Savanna Ocean Tropical forest Interior Southeast Asia Frontier Savanna Ocean Tropical forest Interior Total tropics Frontier Savanna Ocean 288 123 5 921 2966 693 390 64 397 1494 304 16 56 112 2523 1409 529 145 1610 19530 Biological Nitrogen Fixation kg N ha-1 yr-1 25.8 25.2 6.1 -4.5 2.4 -1.2 -5.4 25.7 0.1 4.8 0.3 0.4 25.2 -6.3 0 0.2 2.3 -2.8 25.7 0 1.1 % of BNF 20 19 13 14 6 4 2 8 5 3 13 13 18 9 6 9 6 12 10 2 Net fire N flux kg N ha-1 yr-1 -0.4 25.6 3.4 0 Deposition of fireemitted N kg N ha-1 yr-1 4.2 Fire emissions of Nr kg N ha-1 yr-1 2.5 9.2 5.7 4.7 0.5 25.4 25.9 0.5 0.9 0.9 0.0 0.2 6.8 0.3 1.0 0.3 2.2 4.2 3.1 0 Fire emissions of N2 kg N ha-1 yr-1 2.9 -2.2 -4.4 25.0 0.5 -4.4 25.5 2.0 2.3 1.4 0.8 0 1.9 0.2 3.4 30.4 30.4 30. Same as Table 1 but expressed in units per unit of ecosystem area (kg N ha-1 yr-1).9 3.4 3.3 2.2 1.3 1.1 0.4 9.2 6.3 0.0 1.2 1.4 0.8 0 1.9 1.1 3.2 5.6 0 3.9 1.1 4.4 0.1 % of BNF -3 16 -5 -39 6 -4 1 -21 -10 3 -35 8 -60 -35 6 -10 4 -38 -28 2 45 .5 0 3.4 30.0 0.5 1.8 -14.Table S2.9 2.

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