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  1. Free, publicly-accessible full text available September 1, 2025
  2. Abstract

    Atmospheric nitrogen (N) deposition and climate change are transforming the way N moves through dryland watersheds. For example, N deposition is increasing N export to streams, which may be exacerbated by changes in the magnitude, timing, and intensity of precipitation (i.e., the precipitation regime). While deposition can control the amount of N entering a watershed, the precipitation regime influences rates of internal cycling; when and where soil N, plant roots, and microbes are hydrologically coupled via diffusion; how quickly plants and microbes assimilate N; and rates of denitrification, runoff, and leaching. We used the ecohydrological model RHESSys to investigate (a) how N dynamics differ between N‐limited and N‐saturated conditions in a dryland watershed, and (b) how total precipitation and its intra‐annual intermittency (i.e., the time between storms in a year), interannual intermittency (i.e., the duration of dry months across multiple years), and interannual variability (i.e., variance in the amount of precipitation among years) modify N dynamics and export. Streamflow nitrate (NO3) export was more sensitive to increasing rainfall intermittency (both intra‐annual and interannual) and variability in N‐limited than in N‐saturated model scenarios, particularly when total precipitation was lower—the opposite was true for denitrification which is more sensitive in N‐saturated than N‐limited scenarios. N export and denitrification increased or decreased more with increasing interannual intermittency than with other changes in precipitation amount. This suggests that under future climate change, prolonged droughts that are followed by more intense storms may pose a major threat to water quality in dryland watersheds.

     
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    Free, publicly-accessible full text available April 1, 2025
  3. Abstract

    Wildfires may increase soil emissions of trace nitrogen (N) gases like nitric oxide (NO) and nitrous oxide (N2O) by changing soil physicochemical conditions and altering microbial processes like nitrification and denitrification. When 34 studies were synthesized, we found a significant increase in both NO and N2O emissions up to 1 year post-fire across studies spanning ecosystems globally. However, when fluxes were separated by ecosystem type, we found that individual ecosystem types responded uniquely to fire. Forest soils tended to emit more N2O after fire, but there was no significant effect on NO. Shrubland soils showed significant increases in both NO and N2O emissions after fires; often with extremely large but short-lived NO pulses occurring immediately after fire. Grassland NO emissions increased after fire, but the size of this effect was small relative to shrublands. N2O emissions from burned grasslands were highly variable with no significant effect. To better understand the variation in responses to fire across global ecosystems, more consistent measurements of variables recognized as important controls on soil fluxes of NO and N2O (e.g., N cycling rates, soil water content, pH, and substrate availability) are needed across studies. We also suggest that fire-specific elements like burn severity, microbial community succession, and the presence of char be considered by future studies. Our synthesis suggests that fires can exacerbate ecosystem N loss long after they burn, increasing soil emissions of NO and N2O with implications for ecosystem N loss, climate, and regional air quality as wildfires increase globally.

     
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  4. Soils are the largest source of atmospheric nitrous oxide (N2O), a powerful greenhouse gas. Dry soils rarely harbor anoxic conditions to favor denitrification, the predominant N2O-producing process, yet, among the largest N2O emissions have been measured after wetting summer-dry desert soils, raising the question: Can denitrifiers endure extreme drought and produce N2O immediately after rainfall? Using isotopic and molecular approaches in a California desert, we found that denitrifiers produced N2O within 15 minutes of wetting dry soils (site preference = 12.8 ± 3.92 per mil, δ15Nbulk= 18.6 ± 11.1 per mil). Consistent with this finding, we detected nitrate-reducing transcripts in dry soils and found that inhibiting microbial activity decreased N2O emissions by 59%. Our results suggest that despite extreme environmental conditions—months without precipitation, soil temperatures of ≥40°C, and gravimetric soil water content of <1%—bacterial denitrifiers can account for most of the N2O emitted when dry soils are wetted.

     
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  5. 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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  6. While altered precipitation regimes can greatly impact biodiversity and ecosystem functioning, we lack a comprehensive view of how these impacts are mediated by changes to the seasonality of precipitation (i.e., whether it rains more/less in one season relative to another). Over 2 years, we examined how altered seasonal precipitation influenced annual plant biomass and species richness, Simpson’s diversity, and community composition of annual plant communities in a dryland ecosystem that receives both winter and summer rainfall and has distinct annual plant communities in each season. Using a rainfall exclusion, collection, and distribution system, we excluded precipitation and added water during each season individually and compared responses to control plots which received ambient summer and winter precipitation. In control plots, we found five times greater annual plant biomass, twice as many species, and higher diversity in winter relative to summer. Adding water increased annual plant biomass in summer only, did not change richness or diversity in either summer or winter, and modestly shifted community composition. Excluding precipitation in either season reduced annual plant biomass, richness, and Simpson’s diversity. However, in the second winter season, biomass was higher in the plots where precipitation was excluded in the previous summer seasons suggesting that reduced productivity in the summer may facilitate biomass in the winter. Our results suggest that increased precipitation in summer may have stronger short-term impacts on annual plant biodiversity and ecosystem function relative to increased winter precipitation. In contrast, decreasing precipitation may have ubiquitous negative effects on annual plants across both summer and winter but may lead to increased biomass in the following off-seasons. These patterns suggest that annual plant communities exhibit asymmetries in their community and ecosystem responses to altered seasonal precipitation and that considering the seasonality of precipitation is important for predicting the effects of altered precipitation regimes. 
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  7. 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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