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  1. Abstract Changes in precipitation patterns can influence soil nitrogen (N) cycling by altering the timing and extent of moisture availability. Arid conditions, associated with more frequent and severe droughts, can increase soil inorganic N concentrations when diffusion constraints decouple N sources and sinks, making this N susceptible to loss. Further, shifts in precipitation patterns in one season can “carry over” N into subsequent seasons, known as precipitation legacies. However, how these legacies increase soil N availability through changes in inorganic N production (i.e., gross N mineralization and nitrification) remain unclear. We manipulated the amount and timing of precipitation in a pinyon–juniper dryland by either adding or excluding precipitation during the winter or summer months and then measured the effects on inorganic N production using isotope pool dilutions. Excluding precipitation in the winter wet season—the most severe drought imposed—reduced average (± std. error) gross N mineralization by ~ 2.5 times to 0.3 ± 0.08 µg N g−1soilh−1(p = 0.0463) and gross nitrification by ~ 9.5 times to 0.2 ± 0.05 µg N g−1soilh−1(p < 0.0001) relative to ambient conditions, yet inorganic N production persisted under dry conditions. Because this mineralized N was not subsequently taken up by N sinks, inorganic N accumulated in the soil; soil N concentrations were ~ 6 times greater than ambient conditions (0.5 ± 0.2 µg NO3-N g−1soil) under experimental winter drought. This winter drought also produced a legacy effect the following summer, where gross N mineralization declined by ~ 2 times below ambient (Winter- = 0.4 ± 0.2 µg N g−1soilh−1) even though the drought had ended. Across seasons, precipitation legacy effects reduced gross N mineralization when organic N was depleted, but did not affect gross nitrification, favoring the accumulation of nitrate over ammonium, a more mobile anion susceptible to loss. Precipitation legacies also carried over ammonium from summer to winter, likely due to the buildup of ammonium from dry conditions in the previous summer. Our results highlight how the legacies of past precipitation can increase soil N availability, selecting N species that are vulnerable to loss when ecosystem N sources and sinks decouple. These legacies can favor N loss over retention, contributing to ecosystem N limitation and to overall declining N availability in regions experiencing increasing drought frequency. 
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    Free, publicly-accessible full text available March 9, 2027
  2. ABSTRACT Wildfires can lower soil carbon (C) stocks directly through combustion, but also indirectly during post‐fire recovery if microbial C demands outpace photosynthetic C inputs. However, how much C is respired by soil microorganisms post‐fire may depend on wildfire effects on particulate organic carbon (POC; mostly plant material accessible to microbes) and/or mineral‐associated organic carbon (MAOC; considered C protected by minerals from decomposers), meaning assessment of wildfire impacts on these pools is necessary to predict microbial decomposition rates and, thus, the fate of soil C. Here, we measured POC, MAOC, pyrogenic organic matter C, plant cover, extracellular enzyme activity (EEA), and microbial community abundance and composition 17 days, and 1, 3, and 4 years after the Holy Fire burned 94 km2of fire‐adapted chaparral. The wildfire immediately decreased POC by 50% (from 51 ± 21 to 26 ± 6 g C kg−1) and MAOC by 33% (from 9.3 ± 0.9 to 6.3 ± 0.9 g C kg−1), consistent with MAOC being less vulnerable to loss than POC. POC decreased by another 38% 1 year post‐fire, consistent with increases in microbial abundance and EEA suggesting increased microbial decomposition. Between 1 and 4 years after the fire, cover of the dominant shrub (Arctostaphylos glandulosa) increased from 3.9% ± 1.6% to 16% ± 5.4% (compared to 58% ± 4.6% in unburned plots), marking the end of net soil C losses. Still, soil C did not increase between 1 and 4 years post‐fire, suggesting plant C inputs did not outpace microbial respiration, a finding consistent with isotopically heavier C from microorganisms raising bulk soil δ13C values. As global changes favor increases in wildfire frequency and severity, C losses via combustion and decomposition may outpace plant C inputs during the first 4 years post‐fire in chaparral, slowing the replenishment of soil C stocks. 
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    Free, publicly-accessible full text available August 1, 2026
  3. ABSTRACT Climate change is altering precipitation regimes that control nitrogen (N) cycling in terrestrial ecosystems. In ecosystems exposed to frequent drought, N can accumulate in soils as they dry, stimulating the emission of both nitric oxide (NO; an air pollutant at high concentrations) and nitrous oxide (N2O; a powerful greenhouse gas) when the dry soils wet up. Because changes in both N availability and soil moisture can alter the capacity of nitrifying organisms such as ammonia‐oxidizing bacteria (AOB) and archaea (AOA) to process N and emit N gases, predicting whether shifts in precipitation may alter NO and N2O emissions requires understanding how both AOA and AOB may respond. Thus, we ask: How does altering summer and winter precipitation affect nitrifier‐derived N trace gas emissions in a dryland ecosystem? To answer this question, we manipulated summer and winter precipitation and measured AOA‐ and AOB‐derived N trace gas emissions, AOA and AOB abundance, and soil N concentrations. We found that excluding summer precipitation increased AOB‐derived NO emissions, consistent with the increase in soil N availability, and that increasing summer precipitation amount promoted AOB activity. Excluding precipitation in the winter (the most extreme water limitation we imposed) did not alter nitrifier‐derived NO emissions despite N accumulating in soils. Instead, nitrate that accumulated under drought correlated with high N2O emission via denitrification upon wetting dry soils. Increases in the timing and intensity of precipitation that are forecasted under climate change may, therefore, influence the emission of N gases according to the magnitude and season during which the changes occur. 
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  4. 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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  5. 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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  6. 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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  7. Free, publicly-accessible full text available May 1, 2027
  8. 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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