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  1. Abstract

    Ectomycorrhizal (EM) associations can promote the dominance of tree species in otherwise diverse tropical forests. These EM associations between trees and their fungal mutualists have important consequences for soil organic matter cycling, yet the influence of these EM-associated effects on surrounding microbial communities is not well known, particularly in neotropical forests. We examined fungal and prokaryotic community composition in surface soil samples from mixed arbuscular mycorrhizal (AM) and ectomycorrhizal (EM) stands as well as stands dominated by EM-associatedOreomunnea mexicana(Juglandaceae) in four watersheds differing in soil fertility in the Fortuna Forest Reserve, Panama. We hypothesized that EM-dominated stands would support distinct microbial community assemblages relative to the mixed AM-EM stands due to differences in carbon and nitrogen cycling associated with the dominance of EM trees. We expected that this microbiome selection in EM-dominated stands would lead to lower overall microbial community diversity and turnover, with tighter correspondence between general fungal and prokaryotic communities. We measured fungal and prokaryotic community composition via high-throughput Illumina sequencing of theITS2(fungi) and16SrRNA (prokaryotic) gene regions. We analyzed differences in alpha and beta diversity between forest stands associated with different mycorrhizal types, as well as the relative abundance of fungal functional groups and various microbial taxa. We found that fungal and prokaryotic community composition differed based on stand mycorrhizal type. There was lower prokaryotic diversity and lower relative abundance of fungal saprotrophs and pathogens in EM-dominated than AM-EM mixed stands. However, contrary to our prediction, there was lower homogeneity for fungal communities in EM-dominated stands compared to mixed AM-EM stands. Overall, we demonstrate that EM-dominated tropical forest stands have distinct soil microbiomes relative to surrounding diverse forests, suggesting that EM fungi may filter microbial functional groups in ways that could potentially influence plant performance or ecosystem function.

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  2. Abstract

    Climate change and nitrogen (N) pollution are altering biogeochemical and ecohydrological processes in dryland watersheds, increasing N export, and threatening water quality. While simulation models are useful for projecting how N export will change in the future, most models ignore biogeochemical “hotspots” that develop in drylands as moist microsites in the soil become hydrologically disconnected from plant roots when soils dry out. These hotspots enable N to accumulate over dry periods and rapidly flush to streams when soils wet up. To better project future N export, we developed a framework for representing hotspots using the ecohydrological model RHESSys. We then conducted a series of virtual experiments to understand how uncertainties in model structure and parameters influence N export to streams. Modeled N export was sensitive to three major factors (a) the abundance of hotspots in a watershed: N export increased linearly and then reached an asymptote with increasing hotspot abundance; this occurred because carbon and N inputs eventually became limiting as hotspots displaced vegetation cover, (b) the soil moisture threshold required for subsurface flow from hotspots to reestablish: peak streamflow N export increased and then decreased with an increasing threshold due to tradeoffs between N accumulation and export that occur with increasingly disconnected hotspots, and (c) the rate at which water diffused out of hotspots as soils dried down: N export was generally higher when the rate was slow because more N could accumulate in hotspots over dry periods, and then be flushed more rapidly to streams at the onset of rain. In a case study, we found that when hotspots were modeled explicitly, peak streamflow nitrate export increased by 29%, enabling us to better capture the timing and magnitude of N losses observed in the field. N export further increased in response to interannual precipitation variability, particularly when multiple dry years were followed by a wet year. This modeling framework can improve projections of N export in watersheds where hotspots play an increasingly important role in water quality.

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  3. Soil ammonia (NH3) emissions are seldom included in ecosystem nutrient budgets; however, they may represent substantial pathways for ecosystem nitrogen (N) loss, especially in arid regions where hydrologic N losses are comparatively small. To characterize how multiple factors affect soil NH3 emissions, we measured NH3 losses from 6 dryland sites along a gradient in soil pH, atmospheric N deposition, and rainfall. We also enriched soils with ammonium (NH4+), to determine whether N availability would limit emissions, and measured NH3 emissions with passive samplers in soil chambers following experimental wetting. Because the volatilization of NH3 is sensitive to pH, we hypothesized that NH3 emissions would be higher in more alkaline soils and that they would increase with increasing NH4+ availability. Consistent with this hypothesis, average soil NH3 emissions were positively correlated with average site pH (R2 = 0.88, P = 0.004), ranging between 0.77 ± 0.81 µg N-NH3 m−2 h−1 at the least arid and most acidic site and 24.2 ± 16.0 µg N-NH3 m−2 h−1 at the most arid and alkaline site. Wetting soils while simultaneously adding NH4+ increased NH3 emissions from alkaline and moderately acidic soils (F1,35 = 14.7, P < 0.001), suggesting that high N availability can stimulate NH3 emissions even when pH is less than optimal for NH3 volatilization. Thus, both pH and N availability act as proximate controls over NH3 emissions suggesting that these N losses may limit how much N accumulates in arid ecosystems. 
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  4. Abstract

    Soil drying and wetting cycles can produce pulses of nitric oxide (NO) and nitrous oxide (N2O) emissions with substantial effects on both regional air quality and Earth’s climate. While pulsed production of N emissions is ubiquitous across ecosystems, the processes governing pulse magnitude and timing remain unclear. We studied the processes producing pulsed NO and N2O emissions at two contrasting drylands, desert and chaparral, where despite the hot and dry conditions known to limit biological processes, some of the highest NO and N2O flux rates have been measured. We measured N2O and NO emissions every 30 min for 24 h after wetting soils with isotopically-enriched nitrate and ammonium solutions to determine production pathways and their timing. Nitrate was reduced to N2O within 15 min of wetting, with emissions exceeding 1000 ng N–N2O m−2 s−1and returning to background levels within four hours, but the pulse magnitude did not increase in proportion to the amount of ammonium or nitrate added. In contrast to N2O, NO was emitted over 24 h and increased in proportion to ammonium addition, exceeding 600 ng N–NO m−2 s−1in desert and chaparral soils. Isotope tracers suggest that both ammonia oxidation and nitrate reduction produced NO. Taken together, our measurements demonstrate that nitrate can be reduced within minutes of wetting summer-dry desert soils to produce large N2O emission pulses and that multiple processes contribute to long-lasting NO emissions. These mechanisms represent substantial pathways of ecosystem N loss that also contribute to regional air quality and global climate dynamics.

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  5. Abstract

    Warming‐induced changes in precipitation regimes, coupled with anthropogenically enhanced nitrogen (N) deposition, are likely to increase the prevalence, duration, and magnitude of soil respiration pulses following wetting via interactions among temperature and carbon (C) and N availability. Quantifying the importance of these interactive controls on soil respiration is a key challenge as pulses can be large terrestrial sources of atmospheric carbon dioxide (CO2) over comparatively short timescales. Using an automated sensor system, we measured soil CO2flux dynamics in the Colorado Desert—a system characterized by pronounced transitions from dry‐to‐wet soil conditions—through a multi‐year series of experimental wetting campaigns. Experimental manipulations included combinations of C and N additions across a range of ambient temperatures and across five sites varying in atmospheric N deposition. We found soil CO2pulses following wetting were highly predictable from peak instantaneous CO2flux measurements. CO2pulses consistently increased with temperature, and temperature at time of wetting positively correlated to CO2pulse magnitude. Experimentally adding N along the N deposition gradient generated contrasting pulse responses: adding N increased CO2pulses in low N deposition sites, whereas adding N decreased CO2pulses in high N deposition sites. At a low N deposition site, simultaneous additions of C and N during wetting led to the highest observed soil CO2fluxes reported globally at 299.5 μmol CO2 m−2 s−1. Our results suggest that soils have the capacity to emit high amounts of CO2within small timeframes following infrequent wetting, and pulse sizes reflect a non‐linear combination of soil resource and temperature interactions. Importantly, the largest soil CO2emissions occurred when multiple resources were amended simultaneously in historically resource‐limited desert soils, pointing to regions experiencing simultaneous effects of desertification and urbanization as key locations in future global C balance.

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  6. Abstract

    Climate change is increasing the variability of precipitation, altering the frequency of soil drying‐wetting events and the distribution of seasonal precipitation. These changes in precipitation can alter nitrogen (N) cycling and stimulate nitric oxide (NO) emissions (an air pollutant at high concentrations), which may vary according to legacies of past precipitation and represent a pathway for ecosystem N loss. To identify whether precipitation legacies affect NO emissions, we excluded or added precipitation during the winter growing season in a Pinyon–Juniper dryland and measured in situ NO emissions following experimental wetting. We found that the legacy of both excluding and adding winter precipitation increased NO emissions early in the following summer; cumulative NO emissions from the winter precipitation exclusion plots (2750 ± 972 μg N‐NO m−2) and winter water addition plots (2449 ± 408 μg N‐NO m−2) were higher than control plots (1506 ± 397 μg N‐NO m−2). The increase in NO emissions with previous precipitation exclusion was associated with inorganic N accumulation, while the increase in NO emissions with previous water addition was associated with an upward trend in microbial biomass. Precipitation legacies can accelerate soil NO emissions and may amplify ecosystem N loss in response to more variable precipitation.

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  7. Topographic depressions in upland soils experience anaerobic conditions conducive for iron (Fe) reduction following heavy rainfall. These depressional areas can also accumulate reactive Fe compounds, carbon (C), and nitrate, creating potential hot spots of Fe-mediated carbon dioxide (CO2) and nitrous oxide (N2O) production. While there are multiple mechanisms by which Fe redox reactions can facilitate CO2 and N2O production, it is unclear what their cumulative effect is on CO2 and N2O emissions in depressional soils under dynamic redox. We hypothesized that Fe reduction and oxidation facilitate greater CO2 and N2O emissions in depressional compared to upslope soils in response to flooding. To test this, we amended upslope and depressional soils with Fe(II), Fe(III), or labile C and measured CO2 and N2O emissions in response to flooding. We found that depressional soils have greater Fe reduction potential, which can contribute to soil CO2 emissions during flooded conditions when C is not limiting. Additionally, Fe(II) addition stimulated N2O production, suggesting that chemodenitrification may be an important pathway of N2O production in depressions that accumulate Fe(II). As rainfall intensification results in more frequent flooding of depressional upland soils, Fe-mediated CO2 and N2O production may become increasingly important pathways of soil greenhouse gas emissions. 
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  8. Abstract

    Soil nitrous oxide (N2O) emissions are highly variable in space and time, making it difficult to estimate ecosystem level fluxes of this potent greenhouse gas. While topographic depressions are often evoked as permanent N2O hot spots and rain events are well‐known triggers of N2O hot moments, soil N2O emissions are still poorly predicted. Thus, the objective of this study was to determine how to best use topography and rain events as variables to predict soil N2O emissions at the field scale. We measured soil N2O emissions 11 times over the course of one growing season from 65 locations within an agricultural field exhibiting microtopography. We found that the topographic indices best predicting soil N2O emissions varied by date, with soil properties as consistently poor predictors. Large rain events (>30 mm) led to an N2O hot moment only in the early summer and not in the cool spring or later in the summer when crops were at peak growth and likely had high evapotranspiration rates. In a laboratory experiment, we demonstrated that low heterotrophic respiration rates at cold temperatures slowly depleted soil dissolved O2, thus suppressing denitrification over the 2–3 day timescale typical of field ponding. Our findings show that topographic depressions do not consistently act as N2O hot spots and that rainfall does not consistently trigger N2O hot moments. We assert that the spatiotemporal variation in soil N2O emissions is not always characterized by predictable hot spots or hot moments and that controls on this variation change depending on environmental conditions.

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