Effects of experimental nitrogen deposition on soil organic carbon storage in Southern California drylands.

Atmospheric nitrogen (N) deposition is enriching soils with N across biomes. Soil N enrichment can increase plant productivity and affect microbial activity, thereby increasing soil organic carbon (SOC), but such responses vary across biomes. Drylands cover ~45% of Earth's land area and store ~33% of global SOC contained in the top 1 m of soil. Nitrogen fertilization could, therefore, disproportionately impact carbon (C) cycling, yet whether dryland SOC storage increases with N remains unclear. To understand how N enrichment may change SOC storage, we separated SOC into plant-derived, particulate organic C (POC), and largely microbially derived, mineral-associated organic C (MAOC) at four N deposition experimental sites in Southern California. Theory suggests that N enrichment increases the efficiency by which microbes build MAOC (C stabilization efficiency) if soil pH stays constant. But if soils acidify, a common response to N enrichment, then microbial biomass and enzymatic organic matter decay may decrease, increasing POC but not MAOC. We found that N enrichment had no effect on C fractions except for a decrease in MAOC at one site. Specifically, despite reported increases in plant biomass in three sites and decreases in microbial biomass and extracellular enzyme activities in two sites that acidified, POC did not increase. Furthermore, microbial C use and stabilization efficiency increased in a non-acidified site, but without increasing MAOC. Instead, MAOC decreased by 16% at one of the sites that acidified, likely because it lost 47% of the exchangeable calcium (Ca) relative to controls. Indeed, MAOC was strongly and positively affected by Ca, which directly and, through its positive effect on microbial biomass, explained 58% of variation in MAOC. Long-term effects of N fertilization on dryland SOC storage appear abiotic in nature, such that drylands where Ca-stabilization of SOC is prevalent and soils acidify, are most at risk for significant C loss.

Radiative forcing of methane fluxes offsets net carbon dioxide uptake for a tropical flooded forest.

Wetlands are important sources of methane (CH4 ) and sinks of carbon dioxide (CO2 ). However, little is known about CH4 and CO2 fluxes and dynamics of seasonally flooded tropical forests of South America in relation to local carbon (C) balances and atmospheric exchange. We measured net ecosystem fluxes of CH4 and CO2 in the Pantanal over 2014-2017 using tower-based eddy covariance along with C measurements in soil, biomass and water. Our data indicate that seasonally flooded tropical forests are potentially large sinks for CO2 but strong sources of CH4 , particularly during inundation when reducing conditions in soils increase CH4 production and limit CO2 release. During inundation when soils were anaerobic, the flooded forest emitted 0.11 ± 0.002 g CH4 -C m-2  d-1 and absorbed 1.6 ± 0.2 g CO2 -C m-2  d-1 (mean ± 95% confidence interval for the entire study period). Following the recession of floodwaters, soils rapidly became aerobic and CH4 emissions decreased significantly (0.002 ± 0.001 g CH4 -C m-2  d-1 ) but remained a net source, while the net CO2 flux flipped from being a net sink during anaerobic periods to acting as a source during aerobic periods. CH4 fluxes were 50 times higher in the wet season; DOC was a minor component in the net ecosystem carbon balance. Daily fluxes of CO2 and CH4 were similar in all years for each season, but annual net fluxes varied primarily in relation to flood duration. While the ecosystem was a net C sink on an annual basis (absorbing 218 g C m-2 (as CH4 -C + CO2 -C) in anaerobic phases and emitting 76 g C m-2 in aerobic phases), high CH4 effluxes during the anaerobic flooded phase and modest CH4 effluxes during the aerobic phase indicate that seasonally flooded tropical forests can be a net source of radiative forcings on an annual basis, thus acting as an amplifying feedback on global warming.

Chronic N enrichment and drought alter plant cover and community composition in a Mediterranean-type semi-arid shrubland.

Anthropogenic nitrogen (N) deposition has caused a decline in native plant species and an increase in exotic plant species in many terrestrial ecosystems; however, vegetation change depends on the rate and/or duration of N input, individual species responses, interactions with other resources, and ecosystem properties such as species richness and canopy cover, soil texture, pH, and/or disturbance regime. Native shrub and exotic forb responses to N enrichment were evaluated over a 13-year field experiment in a mature coastal sage scrub (CSS) shrubland of southern California to test the hypothesis that dry-season N input will cause a decline in native shrubs and an increase in exotic annuals. Nitrogen enrichment caused the dominant native shrubs, Artemisia californica and Salvia mellifera, to respond differently, with A. californica initially increasing with N input but declining thereafter and S. mellifera declining consistently over the 13-year-period. Both species exhibited higher canopy dieback during drought conditions, especially in N plots. Brassica nigra, an exotic annual, invaded N plots significantly more than control plots, but only after 10 years of N addition and a prolonged drought, which increased native shrub canopy dieback. These results indicate a possible synergism between N enrichment and drought on native shrub and exotic forb abundance, which would have important implications for plant diversity in semi-arid shrublands of southwest US that are anticipated to experience an increase in anthropogenic N enrichment and the frequency and duration of drought.

Plant hydraulic responses to long-term dry season nitrogen deposition alter drought tolerance in a Mediterranean-type ecosystem.

Anthropogenic nitrogen (N) deposition represents a significant N input for many terrestrial ecosystems. N deposition can affect plants on scales ranging from photosynthesis to community composition, yet few studies have investigated how changes in N availability affect plant water relations. We tested the effects of N addition on plant water relations, hydraulic traits, functional traits, gas exchange, and leaf chemistry in a semi-arid ecosystem in Southern California using long-term experimental plots fertilized with N for over a decade. The dominant species were Artemisia california and Salvia mellifera at Santa Margarita Ecological Reserve and Adenostoma fasciculatum and Ceanothus greggii at Sky Oaks Field Station. All species, except Ceanothus, showed increased leaf N concentration, decreased foliar carbon to N ratio, and increased foliar N isotopic composition with fertilization, indicating that added N was taken up by study species, yet each species had a differing physiological response to long-term N addition. Dry season predawn water potentials were less negative with N addition for all species except Adenostoma, but there were no differences in midday water potentials, or wet season water potentials. Artemisia was particularly responsive, as N addition increased stem hydraulic conductivity, stomatal conductance, and leaf carbon isotopic composition, and decreased wood density. The alteration of water relations and drought resistance parameters with N addition in Artemisia, as well as Adenostoma, Ceanothus, and Salvia, indicate that N deposition can affect the ability of native Southern California shrubs to respond to drought.

Variations in evapotranspiration and climate for an Amazonian semi-deciduous forest over seasonal, annual, and El Niño cycles.

Tropical forests exchange large amounts of water and energy with the atmosphere and are important in controlling regional and global climate; however, climate and evaportranspiration (E) vary significantly across multiple time scales. To better understand temporal patterns in E and climate, we measured the energy balance and meteorology of a semi-deciduous forest in the rainforest-savanna ecotone of northern Mato Grosso, Brazil, over a 7-year period and analyzed regional climate patterns over a 16-year period. Spectral analysis revealed that E and local climate exhibited consistent cycles over annual, seasonal, and weekly time scales. Annual and seasonal cycles were also apparent in the regional monthly rainfall and humidity time series, and a cycle on the order of 3-5.5 years was also apparent in the regional air temperature time series, which is coincident with the average return interval of El Niño. Annual rates of E were significantly affected by the 2002 El Niño. Prior to this event, annual E was on average 1,011 mm/year and accounted for 52% of the annual rainfall, while after, annual E was 931 mm/year and accounted for 42% of the annual rainfall. Our data also suggest that E declined significantly over the 7-year study period while air temperature significantly increased, which was coincident with a long-term, regional warming and drying trend. These results suggest that drought and warming induced by El Niño and/or climate change cause declines in E for semi-deciduous forests of the southeast Amazon Basin.

Estimating of gross primary production in an Amazon-Cerrado transitional forest using MODIS and Landsat imagery.

The acceleration of the anthropogenic activity has increased the atmospheric carbon concentration, which causes changes in regional climate. The Gross Primary Production (GPP) is an important variable in the global carbon cycle studies, since it defines the atmospheric carbon extraction rate from terrestrial ecosystems. The objective of this study was to estimate the GPP of the Amazon-Cerrado Transitional Forest by the Vegetation Photosynthesis Model (VPM) using local meteorological data and remote sensing data from MODIS and Landsat 5 TM reflectance from 2005 to 2008. The GPP was estimated using Normalized Difference Vegetation Index (NDVI) and Enhanced Vegetation Index (EVI) calculated by MODIS and Landsat 5 TM images. The GPP estimates were compared with measurements in a flux tower by eddy covariance. The GPP measured in the tower was consistent with higher values during the wet season and there was a trend to increase from 2005 to 2008. The GPP estimated by VPM showed the same increasing trend observed in measured GPP and had high correlation and Willmott's coefficient and low error metrics in comparison to measured GPP. These results indicated high potential of the Landsat 5 TM images to estimate the GPP of Amazon-Cerrado Transitional Forest by VPM.

Shoot and root biomass production in semi-arid shrublands exposed to long-term experimental N input.

Anthropogenic nitrogen (N) deposition has affected the primary production of terrestrial ecosystems worldwide; however, ecosystem responses often vary over time because of transient responses, interactions between N, precipitation, and/or other nutrients, and changes in plant species composition. Here we report N-induced changes in above- and below-ground standing crop and production over an 11-year period for two semi-arid shrublands, chaparral and coastal sage scrub (CSS), of Southern California. Shrubs were exposed to 50 kgN ha-1 in the fall of each year to simulate the accumulation of dry N deposition, and shoot and root biomass and leaf area index (LAI) were measured every 3 months to assess how biomass production responded to chronic, dry N inputs. N inputs significantly altered above- and below-ground standing crop, production, and LAI; however, N impacts varied over time. For chaparral, N inputs initially increased root production but suppressed shoot production; however, over time biomass partitioning reversed and plants exposed to N had significantly more shoot biomass. In CSS, N inputs caused aboveground production to increase only during wet years, and this interaction between added N and precipitation was due in part to a highly flexible growth response of CSS shrubs to increases in N and water availability and to a shift from slower-growing native shrubs to fast-growing introduced annuals. Together, these results indicate that long-term N inputs will lead to complex, spatially and temporally variable growth responses for these, and similar, Mediterranean-type shrublands.

Experimental dry-season N deposition alters species composition in southern Californian mediterranean-type shrublands.

Nitrogen (N) deposition in heavily polluted southern Californian shrublands is estimated to be 20-45 kg N x ha(-1) x yr(-1), but more exposed locales can receive as much as 145 kg N x ha(-1) x yr(-1). This large anthropogenic N input has the capacity to alter the composition of plant communities. We conducted N-fertilization experiments in chaparral and coastal sage scrub (CSS) stands over a five-year period to test the hypothesis that plant community composition would change in response to dry-season N addition because of an increase in the relative abundance of herbaceous plant species. Our results indicate that dry-season addition of N significantly altered the community composition of CSS but not chaparral. Contrary to our original hypothesis, changes in community composition were due to changes in the relative abundance of dominant shrubs and not herbaceous plant species. Given that community-level responses to changes in resource availability may take years to decades in order to fully materialize, our results suggest that continued dry-season input of N will cause even larger changes in community composition over time. These results have implications for plant species composition and diversity of mediterranean-type shrublands as N deposition increases with population growth and fossil-fuel use.

A scaling approach for quantifying the net CO2 flux of the Kuparuk River Basin, Alaska.

Net CO2 flux measurements conducted during the summer and winter of 1994-96 were scaled in space and time to provide estimates of net CO2 exchange during the 1995-96 (9 May 1995-8 May 1996) annual cycle for the Kuparuk River Basin, a 9200 km2 watershed located in NE Alaska. Net CO2 flux was measured using dynamic chambers and eddy covariance in moist-acidic, nonacidic, wet-sedge, and shrub tundra, which comprise 95% of the terrestrial landscape of the Kuparuk Basin. CO2 flux data were used as input to multivariate models that calculated instantaneous and daily rates of gross primary production (GPP) and whole-ecosystem respiration (R) as a function of meteorology and ecosystem development. Net CO2 flux was scaled up to the Kuparuk Basin using a geographical information system (GIS) consisting of a vegetation map, digital terrain map, dynamic temperature and radiation fields, and the models of GPP and R. Basin-wide estimates of net CO2 exchange for the summer growing season (9 May-5 September 1995) indicate that nonacidic tundra was a net sink of -31.7 ± 21.3 GgC (1 Gg = 109 g), while shrub tundra lost 32.5 ± 6.3 GgC to the atmosphere (negative values denote net ecosystem CO2 uptake). Acidic and wet sedge tundra were in balance, and when integrated for the entire Kuparuk River Basin (including aquatic surfaces), whole basin summer net CO2 exchange was estimated to be in balance (-0.9 ± 50.3 GgC). Autumn to winter (6 September 1995-8 May 1996) estimates of net CO2 flux indicate that acidic, nonacidic, and shrub tundra landforms were all large sources of CO2 to the atmosphere (75.5 ± 8.3, 96.4 ± 11.4, and 43.3 ± 4.7 GgC for acidic, nonacidic, and shrub tundra, respectively). CO2 loss from wet sedge surfaces was not substantially different from zero, but the large losses from the other terrestrial landforms resulted in a whole basin net CO2 loss of 217.2 ± 24.1 GgC during the 1995-96 cold season. When integrated for the 1995-96 annual cycle, acidic (66.4 + 25.25 GgC), nonacidic (64.7 ± 29.2 GgC), and shrub tundra (75.8 ± 8.4 GgC) were substantial net sources of CO2 to the atmosphere, while wet sedge tundra was in balance (0.4 + 0.8 GgC). The Kuparuk River Basin as a whole was estimated to be a net CO2 source of 218.1 ± 60.6 GgC over the 1995-96 annual cycle. Compared to direct measurements of regional net CO2 flux obtained from aircraft-based eddy covariance, the scaling procedure provided realistic estimates of CO2 exchange during the summer growing season. Although winter estimates could not be assessed directly using aircraft measurements of net CO2 exchange, the estimates reported here are comparable to measured values reported in the literature. Thus, we have high confidence in the summer estimates of net CO2 exchange and reasonable confidence in the winter net CO2 flux estimates for terrestrial landforms of the Kuparuk river basin. Although there is larger uncertainty in the aquatic estimates, the small surface area of aquatic surfaces in the Kuparuk river basin (≈ 5%) presumably reduces the potential for this uncertainty to result in large errors in basin-wide CO2 flux estimates.

Post-fire primary production and plant community dynamics in chaparral stands exposed to varying levels of nitrogen deposition.

High levels of atmospheric nitrogen (N) deposition to southern California chaparral shrublands may interact with fire to affect biomass production and plant species composition during secondary succession. To determine the potential interactions between post fire recovery and N deposition we compared rates of aboveground net primary production (ANPP), shrub growth, and the relative abundance of Adenostoma fasciculatum, other sub-dominant shrubs, and herbaceous species of three chaparral stands exposed to different levels of atmospheric N deposition over the first 3 years of post-fire succession. Our data suggest that rates of ANPP (gdw m(-2) month(-1)) and aboveground N storage (gN m(-2) month(-1)) for these chaparral stands were not related to N deposition even though sites exposed to high levels of N deposition had significantly higher rates of shrub growth (gdw plant(-1) month(-1)) and N uptake (gN plant(-1) month(-1)). However, high-N stands were composed of larger shrubs with a lower density, and this trade-off between shrub size and density may explain the low correlation between N deposition and post-fire ANPP. Differences in relative plant species abundance between sites were significantly correlated with N deposition exposure, where stands exposed to high N deposition had a lower relative abundance of A. fasciculatum and a higher relative abundance of other shrub and herbaceous species. While many factors can affect rates and patterns of post-fire recovery, these results suggest that chronic exposure to N deposition may significantly alter plant growth and species composition in successional chaparral stands.

CARBON AND NITROGEN STORAGE IN SOIL AND LITTER OF SOUTHERN CALIFORNIAN SEMI-ARID SHRUBLANDS.

Semi-arid shrublands of southern California, including chaparral and coastal sage, are found in widely varying elevation and microclimatic regimes and are subjected to disturbance such as fire and atmospheric N deposition that have the capacity to alter soil and litter C and N storage. Here we present a case study where soil and litter C and N were measured over 19 months in post-fire chaparral and mature coastal sage stands to assess whether differences in soil and litter C and N between these diverse shrublands could be attributed to differences in elevation, stand age, rainfall, and/or estimated N deposition exposure. Our results indicate that atmospheric N deposition exposure, either alone or in conjunction with other environmental variables (elevation, rainfall, and/or stand age), was the most frequent predictor of the spatial pattern in the soil and litter N and C variables observed. These results are consistent with those reported for high-elevation coniferous forests arrayed along an N deposition gradient in southern California, suggesting that N deposition may affect the soil N and C storage of semiarid shrublands and woodlands in a qualitatively similar manner.