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As the largest biogeochemically active terrestrial reserve of carbon (C), soils have the potential to either mitigate or amplify rates of climate change. Ecosystems with large C stocks and high rates of soil C sequestration, in particular, may have outsized impacts on regional and global C cycles. Montane meadows have large soil C stocks relative to surrounding ecosystems. However, anthropogenic disturbances in many meadows may have altered the balance of C inputs and outputs, potentially converting these soils from net C sinks to net sources of C to the atmosphere. Here, we quantified ecosystem-level C inputs and outputs to estimate the annual net soil C flux from 13 montane meadows spanning a range of conditions throughout the California Sierra Nevada. Our results suggest that meadow soils can be either large net C sinks (577.6 ± 250.5 g C m−2 y−1) or sources of C to the atmosphere (− 391.6 ± 154.2 g C m−2 y−1). Variation in the direction and magnitude of net soil C flux appears to be driven by belowground C inputs. Vegetation species and functional group composition were not associated with the direction of net C flux, but climate and watershed characteristics were. Our results demonstrate that, per unit area, montane meadows hold a greater potential for C sequestration than the surrounding forest. However, legacies of disturbance have converted some meadows to strong net C sources. Accurate quantification of ecosystem-level C fluxes is critical for the development of regional C budgets and achieving global emissions goals.

Background Meadow ecosystems have important ecological functions and support socioeconomic services, yet are subject to multiple stressors that can lead to rapid degradation. In the Sierra Nevada of the western USA, recreational pack stock (horses and mules) use in seasonally wet mountain meadows may lead to soil trampling and meadow degradation, especially when soil water content is high and vegetation is developing. Methods In order to improve the ability to predict meadow vulnerability to soil disturbance from pack stock use, we measured soil resistance (SR), which is an index of vulnerability to trampling disturbance, at two spatial scales using a stratified-random sampling design. We then compared SR to several soil and vegetation explanatory variables that were also measured at the two spatial scales: plant community type (local scale) and topographic gradient class (meadow scale). Results We found that local-scale differences in drivers of SR were contingent on the meadow scale, which is important because multiple spatial scale evaluation of ecological metrics provides a broader understanding of the potential controls on ecological processes than assessments conducted at a single spatial scale. We also found two contrasting explanatory models for drivers of SR at the local scale: (1) soil gravimetric water content effects on soil disaggregation and (2) soil bulk density and root mass influence on soil cohesion. Soil resistance was insufficient to sustain pack stock use without incurring soil deformation in wet plant communities, even when plant cover was maximal during a major drought. Conclusions Our study provides new information on seasonally wet meadow vulnerability to trampling by pack stock animals using multi-scale drivers of SR, including the contrasting roles of soil disaggregation, friction, and cohesion. Our work aims to inform meadow management efforts in the Sierra Nevada and herbaceous ecosystems in similar regions that are subject to seasonal soil saturation and livestock use.

Unlike plants and vertebrates, the ecological preferences, and potential vulnerabilities of soil invertebrates to environmental change, remain poorly understood in terrestrial ecosystems globally. We conducted a cross-biome survey including 83 locations across six continents to advance our understanding of the ecological preferences and vulnerabilities of the diversity of dominant and functionally important soil invertebrate taxa, including nematodes, arachnids and rotifers. The diversity of invertebrates was analyzed through amplicon sequencing. Vegetation and climate drove the diversity and dominant taxa of soil invertebrates. Our results suggest that declines in forest cover and plant diversity, and reductions in plant production associated with increases in aridity, can result in reductions of the diversity of soil invertebrates in a drier and more managed world. We further developed global atlases of the diversity of these important soil invertebrates, which were cross-validated using an independent database. Our study advances the current knowledge of the ecological preferences and vulnerabilities of the diversity and presence of functionally important soil invertebrates in soils from across the globe. This information is fundamental for improving and prioritizing conservation efforts of soil genetic resources and management policies.

Increasing global temperatures are predicted to stimulate soil microbial respiration. The direct and indirect impacts of warming on soil microbes, nevertheless, remain unclear. This is particularly true for understudied subsoil microbes. Here, we show that 4.5 years of whole-profile soil warming in a temperate mixed forest results in altered microbial community composition and metabolism in surface soils, partly due to carbon limitation. However, microbial communities in the subsoil responded differently to warming than in the surface. Throughout the soil profile—but to a greater extent in the subsoil—physiologic and genomic measurements show that phylogenetically different microbes could utilize complex organic compounds, dampening the effect of altered resource availability induced by warming. We find subsoil microbes had 20% lower carbon use efficiencies and 47% lower growth rates compared to surface soils, which constrain microbial communities. Collectively, our results show that unlike in surface soils, elevated microbial respiration in subsoils may continue without microbial community change in the near-term., There is much uncertainty on the response of soil microbial communities to warming, particularly in the subsoil. Here, the authors investigate microbial community and metabolism response to 4.5 years of whole-profile soil warming, finding depth-dependent effects and elevated subsoil microbial respiration.

Sequential chemical extraction has been widely used to study soil phosphorus (P) dynamics and inform nutrient management, but its efficacy for assigning P into biologically meaningful pools remains unknown. Here, we evaluated the accuracy of the modified Hedley extraction scheme using P K-edge X-ray absorption near-edge structure (XANES) spectroscopy for nine carbonate-free soil samples with diverse chemical and mineralogical properties resulting from different degrees of soil development. For most samples, the extraction markedly overestimated the pool size of calcium-bound P (Ca-P, extracted by 1 M HCl) due to (1) P redistribution during the alkaline extractions (0.5 M NaHCO3 and then 0.1 M NaOH), creating new Ca-P via formation of Ca phosphates between NaOH-desorbed phosphate and exchangeable Ca2+ and/or (2) dissolution of poorly crystalline Fe and Al oxides by 1 M HCl, releasing P occluded by these oxides into solution. The first mechanism may occur in soils rich in well-crystallized minerals and exchangeable Ca2+ regardless of the presence or absence of CaCO3, whereas the second mechanism likely operates in soils rich in poorly crystalline Fe and Al minerals. The overestimation of Ca-P simultaneously caused underestimation of the pools extracted by the alkaline solutions. Our findings identify key edaphic parameters that remarkably influenced the extractions, which will strengthen our understanding of soil P dynamics using this widely accepted procedure.

The release of water during snowmelt orchestrates a variety of important belowground biogeochemical processes in seasonally snow-covered ecosystems, including the production and consumption of greenhouse gases (GHGs) by soil microorganisms. Snowmelt timing is advancing rapidly in these ecosystems, but there is still a need to isolate the effects of earlier snowmelt on soil GHG fluxes. For an improved mechanistic understanding of the biogeochemical effects of snowmelt timing during the snow-free period, we manipulated a high-elevation forest that typically receives over two meters of snowfall but little summer precipitation to influence legacy effects of snowmelt timing. We altered snowmelt rates for two years using black sand to accelerate snowmelt and white fabric to postpone snowmelt, thus creating a two- to three-week disparity in snowmelt timing. Soil microclimate and fluxes of carbon dioxide (CO2 ), methane (CH4 ), and nitrous oxide (N2 O) were monitored weekly to monthly during the snow-free period. Microbial abundances were estimated by potential assays near the end of each snow-free period. Although earlier snowmelt caused soil drying, we found no statistically significant effects (p

Soil profiles are rarely homogeneous. Resource availability and microbial abundances typically decrease with soil depth, but microbes found in deeper horizons are still important components of terrestrial ecosystems. By studying 20 soil profiles across the United States, we documented consistent changes in soil bacterial and archaeal communities with depth. Deeper soils harbored communities distinct from those of the more commonly studied surface horizons. Most notably, we found that the candidate phylum Dormibacteraeota (formerly AD3) was often dominant in subsurface soils, and we used genomes from uncultivated members of this group to identify why these taxa are able to thrive in such resource-limited environments. Simply digging deeper into soil can reveal a surprising number of novel microbes with unique adaptations to oligotrophic subsurface conditions., While most bacterial and archaeal taxa living in surface soils remain undescribed, this problem is exacerbated in deeper soils, owing to the unique oligotrophic conditions found in the subsurface. Additionally, previous studies of soil microbiomes have focused almost exclusively on surface soils, even though the microbes living in deeper soils also play critical roles in a wide range of biogeochemical processes. We examined soils collected from 20 distinct profiles across the United States to characterize the bacterial and archaeal communities that live in subsurface soils and to determine whether there are consistent changes in soil microbial communities with depth across a wide range of soil and environmental conditions. We found that bacterial and archaeal diversity generally decreased with depth, as did the degree of similarity of microbial communities to those found in surface horizons. We observed five phyla that consistently increased in relative abundance with depth across our soil profiles: Chloroflexi, Nitrospirae, Euryarchaeota, and candidate phyla GAL15 and Dormibacteraeota (formerly AD3). Leveraging the unusually high abundance of Dormibacteraeota at depth, we assembled genomes representative of this candidate phylum and identified traits that are likely to be beneficial in low-nutrient environments, including the synthesis and storage of carbohydrates, the potential to use carbon monoxide (CO) as a supplemental energy source, and the ability to form spores. Together these attributes likely allow members of the candidate phylum Dormibacteraeota to flourish in deeper soils and provide insight into the survival and growth strategies employed by the microbes that thrive in oligotrophic soil environments.

While individual tree genotypes are known to differ in their impacts on local soil development, the spatial genetic influence of surrounding neighboring trees is largely unknown. We examine the hypothesis that fine root dynamics of a focal tree is based on the genetics of the focal tree as well as the genetics of neighbor trees that together define litter inputs to soils of the focal tree. We used a common garden environment with clonal replicates of individual tree genotypes to analyze fine root production, turnover and allocation with respect to modeled neighborhood: (1) foliar mass, (2) foliar condensed tannins (CT), (3) genetic identity of trees, and (4) genetic dissimilarity of neighbors. In support of our central hypothesis, we found that the presence of genetically dissimilar trees and high leaf CT contributions to the soil predicted increased fine root production. In fact, the modeled effects of neighbors accounted for ~90% of the explanatory weight of all models predicting root production. Nevertheless, the ultimate fate of those roots in soil (turnover) and the balance of fine roots relative to aboveground tree mass were still more predictable based on the leaf traits and genetics of the individual focal trees (explaining 99% of the variation accounted for by models). Our data provide support for a method allowing a comparison of the relative effects of individuals vs. spatial neighborhood effects on soils in a genetic context. Such comparisons are important for placing plant-soil feedbacks in a genetic and evolutionary framework because neighbors can decouple feedbacks between an individual and the surrounding environment.

Identifying the global drivers of soil priming is essential to understanding C cycling in terrestrial ecosystems. We conducted a survey of soils across 86 globally-distributed locations, spanning a wide range of climates, biotic communities, and soil conditions, and evaluated the apparent soil priming effect using 13C-glucose labeling. Here we show that the magnitude of the positive apparent priming effect (increase in CO2 release through accelerated microbial biomass turnover) was negatively associated with SOC content and microbial respiration rates. Our statistical modeling suggests that apparent priming effects tend to be negative in more mesic sites associated with higher SOC contents. In contrast, a single-input of labile C causes positive apparent priming effects in more arid locations with low SOC contents. Our results provide solid evidence that SOC content plays a critical role in regulating apparent priming effects, with important implications for the improvement of C cycling models under global change scenarios., The global ecological predictors of soil priming remain unclear. Here the authors conducted a global survey of soils from 86 global locations using an isotopic approach and find that in more mesic sites with high SOC concentrations, soil priming effects are more likely to be negative.

Aeolian dust deposition is an important phosphorus (P) input to terrestrial ecosystems, but its influence on P dynamics during long-term ecosystem development remains poorly understood. In this study, we characterized P speciation using P K-edge XANES spectroscopy in surface soils (0–15 cm, A horizon) and contemporary aeolian dust collected at each site of a 3000-ky volcanic soil chronosequence in a cool, semi-arid environment. Phosphorus speciation in dust was dominated by calcium-bound P (Ca-P; 54–74%), with 11–23% iron and aluminum-bound P [(Fe + Al)-P] and 7–25% organic P (Po). In soils, Po contributed 1–23% of total P, being greater in older soils; however, the proportions of Ca-P (16–39%) and (Fe + Al)-P (48–82%) fluctuated with increasing weathering over the soil chronosequence. These soil fluctuations resulted from the accumulation and preservation of alkaline aeolian dust during pedogenesis in the semi-arid climate, which significantly increased soil Ca-P while decreasing the total amounts and relative abundances of soil (Fe + Al)-P. We suggest that the effects of an aeolian dust input on soil P transformations are functions of the relative magnitude and chemical composition of the dust input and the soil weathering intensity. For a given source of dust, when the net dust flux is greater than the weathering rate, dust accumulates and thus alters the pattern of P transformations during pedogenesis; otherwise, the dust influence on soil P transformations is negligible. By accurately identifying the chemical nature of P pools, our work highlights the advantage of P K-edge XANES spectroscopy over chemical extractions in examining soil P dynamics, and demonstrates how dust inputs can modify the Walker and Syers model of pedogenic P transformations in semi-arid environments. Overall, this work provides a foundation for understanding how dust influences P cycling during soil and ecosystem development, and indicates that dust inputs and composition, and the soil weathering rate, all must be considered for developing integrated climate-biogeochemical models with predictive power in terrestrial ecosystems.

Soil erosion continuously redistributes soil and associated soil organic matter (SOM) on the Earth's surface, with important implications for biogeochemical cycling of essential elements and terrestrial carbon sequestration. Despite the importance of soil erosion, surprisingly few studies have evaluated the sources of eroded carbon (C). We used natural abundance levels of the stable and radioactive isotopes of C (13C and 14C) and stable isotope of nitrogen (15N) to elucidate the origins of SOM eroded from low-order catchments along the western slopes of the Sierra Nevada of California, USA. Our work was conducted in two relatively undisturbed catchments (low elevation = 1800 m, and high elevation = 2300 m) of the Kings River Experimental Watersheds (KREW) in the Sierra National Forest. Sediment captured in basins at the outlet of each gauged watershed were compared to possible source materials, which included: upland surficial organic horizons (i.e., forest floor) and mineral soils (0–0.6 m) from three landform positions (i.e., crest, backslope, and toeslope), stream bank soils (0–0.6 m), and stream-bed materials (0–0.05 m). We found that most of the organic matter (OM) in the captured sediments was composed of O-horizon material that had high C concentrations. Radiocarbon analyses also showed that the captured OM is composed of modern (post-1950) C, with fraction modern values at or above 1.0. Our results suggest that surface (sheet) erosion, as opposed to channeling through established streams and episodic mass wasting events, is likely the largest source of sediment exported out of these minimally disturbed, headwater catchments. The erosional export of sediment with a high concentration of C, especially in the form of relatively undecomposed litter from the O horizon, suggests that a large fraction of the exported C is likely to be decomposed during or after erosion; hence, it is unlikely that soil erosion acts as a significant net sink for atmospheric CO2 in these low-order, temperate forest catchments.

Shifts in microbial communities driven by anthropogenic nitrogen (N) addition have broad-scale ecological consequences. However, responses of microbial groups to exogenous N supply vary considerably across studies, hindering efforts to predict community changes. We used meta-analytical techniques to explore how amoA gene abundances of ammonia-oxidizing archaea (AOA) and bacteria (AOB) respond to N addition, and found that N addition increased AOA and AOB abundances by an average of 27% and 326%, respectively. Responses of AOB varied by study type, ecosystem, fertilizer type, and soil pH, and were strongest in unmanaged wildland soils and soils fertilized with inorganic N sources. Increases in nitrification potential with N addition significantly correlated with only AOB. Our analyses suggest that elevated N supply enhances soil nitrification potential by increasing AOB populations, and that this effect may be most pronounced in unmanaged wildland soils.

Establishment of native grasses is a primary objective of restoration in Pinus ponderosa var. scopulorum (P. & C. Lawson) forests in the southwestern United States. Interactions among native grasses and soil microorganisms generate feedbacks that influence the achievement of this objective. We examined soil chemical properties and communities of plants and soil microorganisms in clear-cuts and P. ponderosa stands thinned and maintained at low and medium tree densities for over 40 years along with high density (unthinned) stands. Phospholipid fatty acids (PLFA) in soils were analyzed to examine arbuscular mycorrhizal (AM) fungi and microbial communities in the three thinning treatments and the unthinned stands with and without a recent broadcast burn. Additionally, two native bunchgrasses, Festuca arizonica and Muhlenbergia wrightii were grown in containers filled with intact soil cores collected from each field plot to more thoroughly compare the abundance of AM fungi and microbial communities across different stand densities and burn treatments. Tree thinning decreased litter cover and increased the abundance and diversity and altered community composition of both herbaceous vegetation and AM fungi. In the mineral soil layer, the pH, total carbon, nitrogen, phosphorus and PLFA profiles did not differ significantly among the four stand density or burn treatments. Mycorrhizal colonization of the container grown grasses did not significantly differ with tree density or burn treatments; however, F. arizonica roots had a strong trend for decreased colonization when grown in soil from high density (unthinned) tree cover. Soil from the containers with F. arizonica had a greater abundance of AM fungal spores. Furthermore, bacterial community composition varied with grass species. Concentration of biomarkers for bacteria were higher in soil that supported F. arizonica compared to soil in which M. wrightii was grown. Our results indicate that the creation of clear-cut openings in forests may increase the abundance and richness of AM fungal propagules and soil bacterial communities were surprisingly resilient to tree thinning and low-intensity fire treatments. These results suggest managing forests to create clear-cut openings generate conditions that favor understory native grasses and AM fungi that are linked to soil bacterial communities.

Soils are important sources and sinks of three greenhouse gases (GHGs): carbon dioxide (CO2), methane (CH4), and nitrous oxide (N2O). However, it is unknown whether semiarid landscapes are important contributors to global fluxes of these gases, partly because our mechanistic understanding of soil GHG fluxes is largely derived from more humid ecosystems. We designed this study with the objective of identifying the important soil physical and biogeochemical controls on soil GHG fluxes in semiarid soils by observing seasonal changes in soil GHG fluxes across a three million year substrate age gradient in northern Arizona. We also manipulated soil nitrogen (N) and phosphorus availability with 7 years of fertilization and used regression tree analysis to identify drivers of unfertilized and fertilized soil GHG fluxes. Similar to humid ecosystems, soil N2O flux was correlated with changes in N and water availability and soil CO2 efflux was correlated with changes in water availability and temperature. Soil CH4 uptake was greatest in relatively colder and wetter soils. While fertilization had few direct effects on soil CH4 flux, soil nitrate was an important predictor of soil CH4 uptake in unfertilized soils and soil ammonium was an important predictor of soil CH4 uptake in fertilized soil. Like in humid ecosystems, N gas loss via nitrification or denitrification appears to increase with increases in N and water availability during ecosystem development. Our results suggest that, with some exceptions, the drivers of soil GHG fluxes in semiarid ecosystems are often similar to those observed in more humid ecosystems.

Author(s): Coble, AA; Hart, SC; Ketterer, ME; Newman, GS; Kowler, AL | Abstract: ©2015. American Geophysical Union. Without exogenous rock-derived nutrient sources, terrestrial ecosystems may eventually regress or reach a terminal steady state, but the degree to which exogenous nutrient sources buffer or slow to a theoretical terminal steady state remains unclear. We used strontium isotope ratios (87Sr/86Sr) as a tracer and measured 87Sr/86Sr values in aeolian dust, soils, and vegetation across a well-constrained 3Myr semiarid substrate age gradient to determine (1) whether the contribution of atmospheric sources of rock-derived nutrients to soil and vegetation pools varied with substrate age and (2) to determine if the depth of uptake varied with substrate age. We found that aeolian-derived nutrients became increasingly important, contributing as much as 71% to plant-available soil pools and tree (Pinus edulis) growth during the latter stages of ecosystem development in a semiarid climate. The depth of nutrient uptake increased on older substrates, demonstrating that trees in arid regions can acquire nutrients from greater depths as ecosystem development progresses presumably in response to nutrient depletion in the more weathered surface soils. Our results demonstrate that global and regional aeolian transport of nutrients to local ecosystems is a vital process for ecosystem development in arid regions. Furthermore, these aeolian nutrient inputs contribute to deep soil nutrient pools, which become increasingly important for maintaining plant productivity over long time scales.

The current paradigm of plant nutrient limitation during ecosystem development predicts a change from nitrogen (N) limitation when substrates are young to phosphorus (P) limitation when substrates are old. However, there are surprisingly few direct tests of this model. We evaluated this theory experimentally along a three million year semi-arid substrate age gradient using resource additions to intercanopy spaces dominated by the C4 bunchgrass Bouteloua gracilis. Unlike other gradients in subtropical and temperate ecosystems, soil water availability also increases strongly across this semi-arid system due to finer texture with substrate age. We found that aboveground net primary production (ANPP) of B. gracilis was limited by both water and N on the 55 ky substrate; not limited by N, P, or water on the 750 ky substrate; and limited by P alone on the 3000 ky substrate. Notably, measures of foliar nutrient concentration and N:P mass ratios were unable to predict nutrient limitations in these semi-arid systems. In unamended plots, mean ANPP declined dramatically at 3000 ky compared to the younger substrate age sites, presumably due to progressive limitation by P. This decline in ANPP late in ecosystem development is consistent with a reduction in soil total carbon and N storage at this site and provides a mechanism for successional retrogression in ecosystem structure and function. Our results unify biogeochemical theory across disparate ecosystems while illustrating the important water-nutrient interactions in these semi-arid ecosystems to further define the nature of nutrient limitations in terrestrial ecosystems.

Dust provides ecosystem-sustaining nutrients to landscapes underlain by intensively weathered soils. Here we show that dust may also be crucial in montane forest ecosystems, dominating nutrient budgets despite continuous replacement of depleted soils with fresh bedrock via erosion. Strontium and neodymium isotopes in modern dust show that Asian sources contribute 18–45% of dust deposition across our Sierra Nevada, California study sites. The remaining dust originates regionally from the nearby Central Valley. Measured dust fluxes are greater than or equal to modern erosional outputs from hillslopes to channels, and account for 10–20% of estimated millennial-average inputs of bedrock P. Our results demonstrate that exogenic dust can drive the evolution of nutrient budgets in montane ecosystems, with implications for predicting forest response to changes in climate and land use., Dust is an important nutrient source to landscapes, but often the source of dust is poorly constrained. Here, the authors quantify the origin of different dust sources in the Sierra Nevada by analysing dust composition and suggest exogenic dust may drive nutrient budgets in montane ecosystems.

Although herbivores are well known to alter litter inputs and soil nutrient fluxes, their long-term influences on soil development are largely unknown because of the difficulty of detecting and attributing changes in carbon and nutrient pools against large background levels. The early phase of primary succession reduces this signal-to-noise problem, particularly in arid systems where individual plants can form islands of fertility. We used natural variation in tree-resistance to herbivory, and a 15 year herbivore-removal experiment in an Arizona pinon-juniper woodland that was established on cinder soils following a volcanic eruption, to quantify how herbivory shapes the development of soil carbon (C) and nitrogen (N) over 36-54 years (i.e., the ages of the trees used in our study). In this semi-arid ecosystem, trees are widely spaced on the landscape, which allows direct examination of herbivore impacts on the nutrient-poor cinder soils. Although chronic insect herbivory increased annual litterfall N per unit area by 50% in this woodland, it slowed annual tree-level soil C and N accumulation by 111% and 96%, respectively. Despite the reduction in soil C accumulation, short-term litterfall-C inputs and soil C-efflux rates per unit soil surface were not impacted by herbivory. Our results demonstrate that the effects of herbivores on soil C and N fluxes and soil C and N accumulation are not necessarily congruent: herbivores can increase N in litterfall, but over time their impact on plant growth and development can slow soil development. In sum, because herbivores slow tree growth, they slow soil development on the landscape.

In humid ecosystems, the rate of methane (CH4) oxidation by soil-dwelling methane-oxidizing bacteria (MOB) is controlled by soil texture and soil water holding capacity, both of which limit the diffusion of atmospheric CH4 into the soil. However, it remains unclear whether these same mechanisms control CH4 oxidation in more arid soils. This study was designed to measure the proximate controls of potential CH4 oxidation in semiarid soils during different seasons. Using a unique and well-constrained 3-million-year-old semiarid substrate age gradient, we were able to hold state factors constant while exploring the relationship between seasonal potential CH4 oxidation rates and soil texture, soil water holding capacity, and dissolved organic carbon (DOC). We measured unexpectedly higher rates of potential CH4 oxidation in the wet season than the dry season. Although other studies have attributed low CH4 oxidation rates in dry soils to desiccation of MOB, we present several lines of evidence that this may be inaccurate. We found that soil DOC concentration explained CH4 oxidation rates better than soil physical factors that regulate the diffusion of CH4 from the atmosphere into the soil. We show evidence that MOB facultatively incorporated isotopically labeled glucose into their cells, and MOB utilized glucose in a pattern among our study sites that was similar to wet-season CH4 oxidation rates. This evidence suggests that DOC, which is utilized by MOB in other environments with varying effects on CH4 oxidation rates, may be an important regulator of CH4 oxidation rates in semiarid soils. Our collective understanding of the facultative use of DOC by MOB is still in its infancy, but our results suggest it may be an important factor controlling CH4 oxidation in soils from dry ecosystems.

Summary •Soil carbon dioxide (CO2) efflux is a major component of terrestrial carbon (C) cycles; yet, the demonstration of covariation between overstory tree genetic-based traits and soil C flux remains a major frontier in understanding biological controls over soil C. •Here, we used a common garden with two native tree species, Populus fremontii and P. angustifolia, and their naturally occurring hybrids to test the predictability of belowground C fluxes on the basis of taxonomic identity and genetic marker composition of replicated clones of individual genotypes. •Three patterns emerged: soil CO2 efflux and ratios of belowground flux to aboveground productivity differ by as much as 50–150% as a result of differences in clone identity and cross type; on the basis of Mantel tests of molecular marker matrices, we found that c. 30% of this variation was genetically based, in which genetically similar trees support more similar soil CO2 efflux under their canopies than do genetically dissimilar trees; and the patterns detected in an experimental garden match observations in the wild, and seem to be unrelated to measured abiotic factors. •Our findings suggest that the genetic makeup of the plants growing on soil has a significant influence on the release of C from soils to the atmosphere.

Background and aims Soils can act as agents of natural selection, causing differential fitness among genotypes and/or families of the same plant species, especially when soils have extreme physical or chemical properties. More subtle changes in soils, such as variation in microbial communities, may also act as agents of selection. We hypothesized that variation in soil properties within a single river drainage can be a selective gradient, driving local adaptation in plants. Methods Using seeds collected from individual genotypes of Populus angustifolia James and soils collected from underneath the same trees, we use a reciprocal transplant design to test whether seedlings would be locally adapted to their parental soil type. Results We found three patterns: 1. Soils from beneath individual genotypes varied in pH, soil texture, nutrient content, microbial biomass and the physiological status of microorganisms. 2. Seedlings grown in local soils experienced 2.5-fold greater survival than seedlings planted in non-local soils. 3. Using a composite of height, number of leaves and leaf area to measure plant growth, seedlings grew ∼17.5% larger in their local soil than in non-local soil. Conclusions These data support the hypothesis that variation in soils across subtle gradients can act as an important selective agent, causing differential fitness and local adaptation in plants.

In order to study controls on metabolic processes in soils, we determined the dynamics of 13 CO2 production from two position-specific 13 C-labeled pyruvate isotopologues in the presence and absence of glucose, succinate, pine, and legume leaf litter, and under anaerobic conditions. We also compared 13 CO2 production in soils along a semiarid substrate age gradient in Arizona. We observed that the C from the carboxyl group (C1) of pyruvate was lost as CO2 much faster than its other C atoms (C2,3). Addition of glucose, pine and legume leaf litter reduced the ratio between 13 CO2 production from 1- 13 C pyruvate and 2,3- 13 C pyruvate (C1/C2,3 ratio), whereas anaerobic conditions increased this ratio. Young volcanic soils exhibited a lower C1/C2,3 ratio than older volcanic soils. We interpret a low C1/C2,3 ratio as an indication of increased Krebs cycle activity in response to carbon inputs, while the higher ratio implies a reduced Krebs cycle activity in response to anaerobic conditions. Succinate, a gluconeogenic substrate, reduced 13 CO2 production from pyruvate to near zero, likely reflecting increased carbohydrate biosynthesis from Krebs cycle intermediates. The difference in 13 CO2 production rate from pyruvate isotopologues disappeared 4e5 days after pyruvate addition, indicating that C positions were scrambled by ongoing soil microbial transformations. This work demonstrates that metabolic tracers such as pyruvate can be used to determine qualitative aspects of C flux patterns through metabolic pathways of soil microbial communities. Understanding the controls over metabolic processes in soil may improve our understanding of soil C cycling processes.

Severe wildfire may cause long-term changes in the soil-atmosphere exchange of carbon dioxide and methane, two gases known to force atmospheric warming. We examined the effect of a severe wildfire 10 years after burning to determine decadal-scale changes in soil gas fluxes following fire, and explored mechanisms responsible for these dynamics. We compared soil carbon dioxide efflux, methane uptake, soil temperature, soil water content, soil O horizon mass, fine root mass, and microbial biomass between a burned site and an unburned site that had similar stand conditions to the burned site before the fire. Compared to the unburned site, soil carbon dioxide efflux was 40% lower and methane uptake was 49% higher at the burned site over the 427-day measurement period. Soil O horizon mass, microbial biomass, fine root mass, and surface soil water content were lower at the burned site than the unburned site, but soil temperature was higher. A regression model showed soil carbon dioxide efflux was more sensitive to changes in soil temperature at the burned site than the unburned site. The relative importance of methane uptake to carbon dioxide efflux was higher at the burned site than the unburned site, but methane uptake compensated for only 1.5% of the warming potential of soil carbon dioxide efflux at the burned site. Our results suggest there was less carbon available at the burned site for respiration by plants and microbes, and the loss of the soil O horizon increased methane uptake in soil at the burned site.

Soils are one of the first selective environments a seed experiences and yet little is known about the evolutionary consequences of plant-soil feedbacks. We have previously found that plant phytochemical traits in a model system, Populus spp., influence rates of leaf litter decay, soil microbial communities and rates of soil net nitrogen mineralization. Utilizing this natural variation in plant-soil linkages we examined two related hypotheses: (1) Populus angustifolia seedlings are locally adapted to their native soils; and (2) Soils act as agents of selection, differentially affecting seedling survival and the heritability of plant traits. We conducted a greenhouse experiment by planting seedlings from 20 randomly collected P. angustifolia genetic families in soils conditioned by various Populus species and measured subsequent survival and performance. Even though P. angustifolia soils are less fertile overall, P. angustifolia seedlings grown in these soils were twice as likely to survive, grew 24% taller, had 27% more leaves, and 29% greater above-ground biomass than P. angustifolia seedlings grown in non-native P. fremontii or hybrid soils. Increased survival resulted in higher trait variation among seedlings in native soils compared to seedlings grown in non-native soils. Soil microbial biomass varied significantly across soil environments which could explain more of the variation in seedling performance than soil texture, pH, or nutrient availability, suggesting strong microbial interactions and feedbacks between plants, soils, and associated microorganisms. Overall, these data suggest that a “home-field advantage” or a positive plant soil feedback helps maintain genetic variance in P. angustifolia seedlings.

The Walker and Syers model of phosphorus (P) transformations during pedogenesis is widely accepted for the development of humid ecosystems, but long-term P dynamics of more arid ecosystems remain poorly understood. We tested the Walker and Syers model in semiarid piñon-juniper woodlands by measuring soil P fractions under tree canopies and in intercanopy spaces along a well-constrained, approximately 3000 ka (1 ka = 1000 years) volcanic substrate age gradient in northern Arizona, USA. The various pools of soil P behaved largely as predicted; total soil P and primary mineral P declined consistently with substrate age, labile inorganic P increased early in soil development and then declined at later stages, and organic phosphorus increased consistently across the chronosequence. Within each site, soils under tree canopies tended to have higher concentrations of labile and intermediately available P fractions compared to intercanopy soils. However, the degree of spatial heterogeneity conferred by tree islands was moderated by the stage of soil development. In contrast, tree islands had no influence on within-site distribution of more recalcitrant soil P pools, which appear to be controlled solely by the stage of pedogenesis. Coincident with declines in total P, primary mineral P, and labile inorganic P, we found that phosphatase enzyme activity increased with substrate age; a result consistent with greater ecosystem-level P demand on older, more highly weathered substrates. Our results suggest that, compared to humid climates, reduced inputs of water, energy, and acidity to semiarid ecosystems slow the rate of change in P fractions during pedogenesis, but the overall pattern remains consistent with the Walker and Syers model. Furthermore, our data imply that pedogenic change may be an important factor controlling the spatial distribution of labile P pools in semiarid ecosystems. Taken together, these data should both broaden and unify terrestrial ecosystem development theory.

Understanding covariance of plant genetics and soil processes may improve our understanding the role of plant genetics in structuring soils and ecosystem function across landscapes. We measured soil nitrogen (N) and phosphorus (P) availability using ion exchange resin bags within three river drainages across Utah and Arizona, USA. The three drainages spanned more than 1,000 km in distance, 8° of latitude, and varying climatic regimes, but were similarly dominated by stands of Populus fremontii (S. Watts), P. angustifolia (James), or natural hybrids between the two species. Soil N availability was consistently greater in P. fremontii stands compared to P. angustifolia stands, and hybrid stands were intermediate. However, we found that the influence of overstory type on soil P availability depended on the river drainage. Our study suggests that, even with a near doubling of mean soil N availability across these drainages, the relative genetic-based effects of the dominant plant on N availability remained consistent. These results expand upon previous work by: 1) providing evidence for linkages between plant genetic factors and ecosystem function across geographic scales; and 2) indicating that plant genetic-based effects on nutrient dynamics in a given ecosystem may differ among nutrients (e.g., N vs. P).

Soil microbial organisms are central to carbon (C) and nitrogen (N) transformations in soils, yet not much is known about the stable isotope composition of these essential regulators of element cycles. We investigated the relationship between C and N availability and stable C and N isotope composition of soil microbial biomass across a three million year old semiarid substrate age gradient in northern Arizona. The δ15N of soil microbial biomass was on average 7.2‰ higher than that of soil total N for all substrate ages and 1.6‰ higher than that of extractable N, but not significantly different for the youngest and oldest sites. Microbial 15N enrichment relative to soil extractable and total N was low at the youngest site, increased to a maximum after 55,000 years, and then decreased slightly with age. The degree of 15N enrichment of microbial biomass correlated negatively with the C:N mass ratio of the soil extractable pool. The δ13C signature of soil microbial biomass was 1.4‰ and 4.6‰ enriched relative to that of soil total and extractable pools respectively and showed significant differences between sites. However, microbial 13C enrichment was unrelated to measures of C and N availability. Our results confirm that 15N, but not 13C enrichment of soil microbial biomass reflects changes in C and N availability and N processing during long-term ecosystem development.

We compared energy fluxes between a site converted from ponderosa pine (Pinus ponderosa) forest to sparse grassland by a severe wildfire 10 years ago and a nearby, unburned forest. We used eddy covariance and associated instruments to measure total radiation, net radiation, albedo, and fluxes of energy into latent heat, sensible heat, and the soil. Total radiation, vapor pressure deficit, and air temperature were similar for each site. Compared to the unburned site, net radiation efficiency (net radiation/total radiation) was 30% lower and albedo 30% higher at the burned site. The magnitude of sensible and latent heats varied seasonally at both sites. Sensible heat was the major component of the energy balance in cold or dry seasons, whereas latent heat was the major component in the warm and wet season. Soil heat flux was the smallest in magnitude of the measured energy fluxes. Compared with the unburned forest, the burn-created grassland generally had lower sensible and latent heats, but greater soil heat flux for both soil cooling in winter and warming in summer. The grassland had similar maximum air temperature as the forest, and warmer surface soil temperature during the summer. Thus, the lower albedo and greater sensible heat of the forest did not produce a warmer site compared with the grassland, apparently because of the cooling effect of greater latent heat in the forest. Our results suggest only small changes in site air temperature, but larger changes in site surface soil temperature by shifts from forest to grassland caused by severe fire in northern Arizona ponderosa pine forests.

As atmospheric CO 2 increases, ecosystem carbon sequestration will largely depend on how global changes in climate will alter the balance between net primary production and decomposition. The response of primary production to climatic change has been examined using well-validated mechanistic models, but the same is not true for decomposition, a primary source of atmospheric CO 2 . We used the Long-term Intersite Decomposition Experiment Team (LIDET) dataset and model-selection techniques to choose and parameterize a model that describes global patterns of litter decomposition. Mass loss was best represented by a three-pool negative exponential model, with a rapidly decomposing labile pool, an intermediate pool representing cellulose, and a recalcitrant pool. The initial litter lignin/nitrogen ratio defined the size of labile and intermediate pools. Lignin content determined the size of the recalcitrant pool. The decomposition rate of all pools was modified by climate, but the intermediate pool's decomposition rate was also controlled by relative amounts of litter cellulose and lignin (indicative of lignin-encrusted cellulose). The effect of climate on decomposition was best represented by a composite variable that multiplied a water-stress function by the Lloyd and Taylor variable Q 10 temperature function. Although our model explained nearly 70% of the variation in LIDET data, we observed systematic deviations from model predictions. Below- and aboveground material decomposed at notably different rates, depending on the decomposition stage. Decomposition in certain ecosystem-specific environmental conditions was not well represented by our model; this included roots in very wet and cold soils, and aboveground litter in N-rich and arid sites. Despite these limitations, our model may still be extremely useful for global modeling efforts, because it accurately (R 2 = 0.6804) described general patterns of long-term global decomposition for a wide array of litter types, using relatively minimal climatic and litter quality data.

Summary 1. Ecological restoration often involves returning ecosystem structure to some predisturbance reference state, but ecosystem function must also recover if restoration efforts are to be self-sustaining over the long term. In the south-western United States, ponderosa pine forest structure was altered by disruption of the fire regime following Euro-American settlement. Forest structure is now being restored to presettlement conditions through the application of thinning and burning treatments. However, the effects of these treatments on below-ground ecosystem processes remain unclear. 2. We conducted a water and nitrogen (N) addition experiment in adjacent restored and unrestored ponderosa pine stands and compared soil CO 2 efflux in response to these treatments over a 13-month period. Our goals were to (i) quantify water and N limitation to below-ground carbon (C) cycling in contemporary high-density ponderosa pine forests; and (ii) determine if restoration alleviates water and N limitations. 3. Restoration thinning and burning increased soil CO 2 efflux, along with surface soil water content, temperature and herbaceous fine root biomass, while total fine root biomass decreased as a result of restoration. 4. Water and N additions increased C flux from soils to a similar degree in both restored and unrestored ponderosa pine stands, but the increase was relatively small when compared to that stimulated by restoration. 5. Synthesis and applications. An understanding of how ecosystem processes respond to treatments designed to restore ecosystem structure is critical in ensuring the long-term success of restoration efforts. Here we show that, although water and N stimulate C flux from soils in these semi-arid forests, restoration treatments have a much greater effect on soil C balance than increased water and N availability by themselves. This suggests that increased quality of C inputs from a recovering understorey herbaceous community is a key component of restoring ecosystem function (e.g. below-ground C cycling) in south-western ponderosa pine forests.

We report the first simultaneous measurements of δ 15 N and δ 13 C of DNA extracted from surface soils. The isotopic composition of DNA differed significantly among nine different soils. The δ 13 C and δ 15 N of DNA was correlated with δ 13 C and δ 15 N of soil, respectively, suggesting that the isotopic composition of DNA is strongly influenced by the isotopic composition of soil organic matter. However, in all samples DNA was enriched in 13 C relative to soil, indicating microorganisms fractionated C during assimilation or preferentially used 13 C enriched substrates. Enrichment of DNA in 15 N relative to soil was not consistently observed, but there were significant differences between δ 15 N of DNA and δ 15 N of soil for three different sites, suggesting microorganisms are fractionating N or preferentially using N substrates at different rates across these contrasting ecosystems. There was a strong linear correlation between δ 15 N of DNA and δ 15 N of the microbial biomass, which indicated DNA was depleted in 15 N relative to the microbial biomass by approximately 3.4‰. Our results show that accurate and precise isotopic measurements of C and N in DNA extracted from the soil are feasible, and that these analyses may provide powerful tools for elucidating C and N cycling processes through soil microorganisms.

© 2015 Carey, Beman, Eviner, Malmstrom and Hart. Global and regional environmental changes often co-occur, creating complex gradients of disturbance on the landscape. Soil microbial communities are an important component of ecosystem response to environmental change, yet little is known about how microbial structure and function respond to multiple disturbances, or whether multiple environmental changes lead to unanticipated interactive effects. Our study used experimental semi-arid grassland plots in a Mediterranean-climate to determine how soil microbial communities in a seasonally variable ecosystem respond to one, two, or three simultaneous environmental changes: exotic plant invasion, plant invasion + vegetation clipping (to simulate common management practices like mowing or livestock grazing), plant invasion + nitrogen (N) fertilization, and plant invasion + clipping + N fertilization. We examined microbial community structure 5-6 years after plot establishment via sequencing of >1 million 16S rRNA genes. Abiotic soil properties (soil moisture, temperature, pH, and inorganic N) and microbial functioning (nitrification and denitrification potentials) were also measured and showed treatment-induced shifts, including altered NO3-availability, temperature, and nitrification potential. Despite these changes, bacterial and archaeal communities showed little variation in composition and diversity across treatments. Even communities in plots exposed to three interacting environmental changes were similar to those in restored native grassland plots. Historical exposure to large seasonal and inter-annual variations in key soil properties, in addition to prior site cultivation, may select for a functionally plastic or largely dormant microbial community, resulting in a microbial community that is structurally robust to single and multiple environmental changes.

Stable isotope analysis is a powerful tool in the study of soil organic matter formation. It is often observed that more decomposed soil organic matter is 13 C, and especially 15 N-enriched relative to fresh litter and recent organic matter. We investigated whether this shift in isotope composition relates to the isotope composition of the microbial biomass, an important source for soil organic matter. We developed a new approach to determine the natural abundance C and N isotope composition of the microbial biomass across a broad range of soil types, vegetation, and climates. We found consistently that the soil microbial biomass was 15 N-enriched relative to the total (3.2 %) and extractable N pools (3.7 %), and 13 C-enriched relative to the extractable C pool (2.5 %). The microbial biomass was also 13 C-enriched relative to total C for soils that exhibited a C3-plant signature (1.6 %), but 13 C-depleted for soils with a C4 signature (� 1.1 %). The latter was probably associated with an increase of annual C3 forbs in C4 grasslands after an extreme drought. These findings are in agreement with the proposed contribution of microbial products to the stabilized soil organic matter and may help explain the shift in isotope composition during soil organic matter formation. r 2006 Elsevier Ltd. All rights reserved.

Soils that are physically disturbed are often reported to show net nitrification and NO3 loss. To investigate the response of soil N cycling rates to soil mixing, we assayed gross rates of mineralization, nitrification, NH4 consumption, and NO3 consumption in a suite of soils from eleven woody plant communities in Oregon, New Mexico, and Utah. Results suggest that the com- mon response of net NO3 flux from disturbed soils is not a straightforward response of increased gross nitrification, but instead may be due to the balance of several factors. While mineralization and NH4 assimilation were higher in mixed than intact cores, NO3 consumption declined. Mean net nitrification was 0.12 mg N kg -1 d -1 in dis- turbed cores, which was significantly higher than in intact cores (-0.19 mg N kg -1 d -1 ). However, higher net nitrification rates in disturbed soils were due to the suppression of NO3 consumption, rather than an increase in nitrification. Our results suggest that at least in the short term, disturbance may significantly increase NO3 flux at the eco

Relatively little research has been conducted on how climate change may affect the structure and function of arid to semiarid ecosystems of the American Southwest. Along the slopes of the San Francisco Peaks of Arizona, USA, I transferred intact soil cores from a spruce-fir to a ponderosa pine forest 730m lower in elevation to assess the potential impacts of climate change on soil N cycling and trace gas fluxes. The low-elevation site has a mean annual soil temperature about 2.51C higher than the high-elevation site. Net rates of N transformations and trace gas fluxes were measured in high-elevation soil cores incubated in situ and soil cores transferred to the low-elevation site. Over a 13-month period, volumetric soil water content was similar in transferred soil cores relative to soil cores incubated in situ. Net N mineralization and nitrification increased over 80% in transferred soil cores compared with in situ soil cores. Soil transfer significantly increased net CO2 efflux (120%) and net CH4 consumption (90%) relative to fluxes of these gases from soil cores incubated in situ. Soil net N2O fluxes were relatively low and were not generally altered by soil transfer. Although the soil microbial biomass as a whole decreased in transferred soil cores compared with in situ soil cores after the incubation period, active bacterial biomass increased. Transferring soil cores from the low-elevation to the high-elevation site (i.e. simulated global cooling) commonly, but not consistently, resulted in the opposite effects on soil pools and processes. In general, soil containment (root trenching) did not significantly affect soil measurements. My results suggest that small increases in mean annual temperature can have large impacts on soil N cycling, soil‐atmosphere trace gas exchanges, and soil microbial communities even in ecosystems where water availability is a major limiting resource.

The availability of C and N to the soil microbial biomass is an important determinant of the rates of soil N transformations. Here, we present evidence that changes in C and N availability affect the 15 N natural abundance of the microbial biomass relative to other soil N pools. We analysed the 15 N natural abundance signature of the chloroform-labile, extractable, NO 3 - , NH 4 + and soil total N pools across a cattle manure gradient associated with a water reservoir in semiarid, high-desert grassland. High levels of C and N in soil total, extractable, NO 3 - , NH 4 + and chloroform-labile fractions were found close to the reservoir. The δ 15 N value of chloroform-labile N was similar to that of extractable (organic + inorganic) N and NO 3 - at greater C availability close to the reservoir, but was 15 N-enriched relative to these N-pools at lesser C availability farther away. Possible mechanisms for this variable 15 N-enrichment include isotope fractionation during N assimilation and dissimilation, and changes in substrate use from a less to a more 15 N-enriched substrate with decreasing C availability.

Ecologists have tried to link plant species composition and ecosystem properties since the inception of the ecosystem concept in ecology. Many have observed that biological communities could feed back to, and not simply result from, soil properties. But which group of organisms, plants or microorganisms, drive those feedback systems? Recent research asserts that soil microorganisms preclude plant species feedback to soil nitrogen (N) transformations due to strong microbial control of soil N cycling. It has been well documented that litter properties influence soil N cycling. In this review, we stress that under many circumstances plant species exert a major influence over soil N cycling rates via unique N attainment strategies, thus influencing soil N availability and their own fitness. We offer two testable mechanisms by which plants impart active control on the N cycle and thereby allow for plant-litter-soil-plant feedback. Finally, we describe the characteristics of plants and ecosystems that are most likely to exhibit feedback.

Nitrogen-fixing tree species have been shown to improve site fertility and increase N transformation rates, but the influence of N-fixing plants on the soil microbial community as a whole is largely unknown. We used patterns of individual carbon-source utilization and enzyme activities to assess the relative effects of N-fixing red alder on the soil microbial community in three adjacent stands (pure conifer, mixed alder-conifer, and pure alder) of a highly productive coastal Oregon forest where the density of red alder has been experimentally manipulated for over 65 years. Two major patterns were revealed: (1) bacterial and fungal carbon-source utilization patterns in soil from pure conifer stands were significantly different from both pure alder soils and mixed conifer-alder soils, while there was no difference in substrate utilization patterns between soils from the mixed alder-conifer and pure alder stands; and (2) the activities of nine extracellular enzymes involved in ligno-cellulose degradation and the mineralization of organic nitrogen, phosphate, and sulfate compounds were all significantly greater in pure alder soils compared to either pure conifer or mixed conifer-alder soils. Our results show that, in addition to an overall increase in soil fertility, microbial biomass, and microbial activity, the presence of N-fixing red alder significantly alters the physiological profile of the microbial community-even in an ecosystem already of high N status.

species. The risk of stand-replacing fire has also greatly increased due to heavy fuel loading and high tree den- control duringeither the dryor wet periodsof the growingseason. Soil macroscale impacts of ecological restoration in this re- respiration measured in situ was significantly higher in the restoration gion (e.g., Stone et al., 1999; Bailey and Covington, 2002), treatments than in the control only during the dry period, while soil enzyme activities were generally higher in the composite restoration little emphasis has been given to possible impacts on the treatmentthaninthethinningrestorationorcontroltreatmentsduring soil microflora. Soil microorganisms play fundamental the wet period. Community-level physiological profiles suggested dif- roles in many ecosystem processes, including decompo- ferences in the physiological capacities of bacteria and fungi in the sition and nutrient cycling, and affect many important composite restoration treatment compared with the other treatments. soil hydrological and chemical properties (Gallardo and We also compared microbial characteristics under different canopy Schlesinger, 1994; Hart et al., 2005). Hence, changes in types to evaluate the impacts of pine invasion and establishment in grass the soil microbial community may lead to changes in openings on soil microorganisms. Soil respiration rates (dry period only) the structure and function of the overall ecosystem, and and enzyme activities (wet period only) were higher in grass openings ultimately determine ecosystem sustainability (Bossio than under presettlement trees, with intermediate values found under postsettlement pines that have invaded grass areas. Taken together, and Scow, 1995). our results suggest that restoration treatments have long-term impacts Soils disturbed by tree removal often have reduced on the soil microflora in these forests. microbial diversity compared with undisturbed areas (Atlas et al., 1991; Arunachalam et al., 1999; Byrd et al., 2000). Furthermore, prescribed fire has been shown to

We investigated the influence of fumigation conditions of the chloroform fumigation‐ extraction method on the release of extractable carbon (C) and nitrogen (N) from organic and mineral soil horizons of 11 mature forests. Soil samples were fumigated with chloroform vapor for 1, 3, 5, or 10 d at two different water contents: field moist or field capacity (233 kPa matric potential). We found that for approximately half our soils, 0.5 M K2SO4-extractable N and C reached a maximum after 1 d of fumigation. The effect of soil wetting on C and N flushes from O horizons was variable but in mineral soils, increasing the soil water content generally resulted in more extractable C and N following fumigation. For the majority of soils assessed, increasing the soil water content did not change the fumigation time necessary to generate the maximum C or N flush. Observed changes in the C/N ratios of the fumigation flush with changes in fumigation time and the sensitivity of this C/N ratio to changes in soil water content suggest that different fumigation conditions may result in different organic pools being extracted. These effects appear to be extremely soil specific. We recommend that the effects of fumigation time and water content be evaluated before the C and N flushes from the fumigation extraction method are used to assess differences in microbial C and N among contrasting forest soils. q 2002 Elsevier Science Ltd. All rights reserved.

Nitric oxide (NO) is a relatively short-lived trace gas that reacts with oxygen in the troposphere to produce the air pollutant ozone. It also reacts with water vapor to form nitric and nitrous acids, which acidify precipitation and increase N deposition. Models currently used to predict soil NO fluxes are based on the assumption that NO flux is proportional to the gross rate of nitrification or N mineralization; however, this assumption has not been tested because of the difficulty in measuring gross N-cycling rates in situ. We measured soil NO fluxes, gross and net N-cycling rates, and a variety of other soil char- acteristics in the forest floor and intact soil cores at nine undisturbed forest and rangeland ecosystems of New Mexico, Utah, and Oregon, USA, to determine which soil variables were most closely related to soil NO flux. Soil NO fluxes ranged from a low of 0.02 ng N·m 22 ·s 21 , prior to wetting in a western hemlock-sitka spruce forest on the Oregon coast, to a high of 6.74 ng N·m 22 ·s 21 , one hour after soil wetting in a juniper woodland of central Oregon. In contrast to our expectations, neither gross nitrification nor gross mineralization was correlated with soil NO flux. Fluxes were positively correlated with net rates of min- eralization and nitrification, soil NO3 2 concentrations, bulk density, and pH, and negatively correlated with gross rates of NO3 2 consumption in the forest floor, soil organic carbon (SOC), soil C:N, and soil water content. Principal-component analysis showed that NO flux after water addition (2 cm of water) had a strong negative correlation with microbial demand for N (as indicated by net mineralization, net nitrification, SOC, and C:N). Our results suggest that, even in well-drained soils, NO efflux is limited more by NO con- sumption than by NO production. As a result, models utilizing the more easily measured net rates, rather than gross rates, may be better predictors of soil NO fluxes across a range of ecosystems.

Roughly one-third of the Earth's land surface is seasonally covered by snow. In many of these ecosystems, the spring snowpack is melting earlier due to climatic warming and atmospheric dust deposition, which could greatly modify soil water resources during the growing season. Though snowmelt timing is known to influence soil water availability during summer, there is little known about the depth of the effects and how long the effects persist. We therefore manipulated the timing of seasonal snowmelt in a high-elevation mixed-conifer forest in a Mediterranean climate during consecutive wet and dry years. The snow-all-gone (SAG) date was advanced by 6 days in the wet year and 3 days in the dry year using black sand to reduce the snow surface albedo. To maximize variation in snowmelt timing, we also postponed the SAG date by 8 days in the wet year and 16 days in the dry year using white fabric to shade the snowpack from solar radiation. We found that deeper soil water (30-60 cm) did not show a statistically significant response to snowmelt timing. Shallow soil water (0-30 cm), however, responded strongly to snowmelt timing. The drying effect of accelerated snowmelt lasted 2 months in the 0-15 cm depth and at least 4 months in the 15-30 cm depth. Therefore, the legacy of snowmelt timing on soil moisture can persist through dry periods, and continued earlier snowmelt due to climatic warming and windblown dust could reduce near-surface water storage and availability to plants and soil biota. Key Points The hydrological signal of snowmelt timing was strongest in shallow soil Effects of snowmelt timing on soil moisture lasted 2-4 months Advancing snowmelt timing by 2-3 weeks depleted shallow soil water by one third © 2014. American Geophysical Union. All Rights Reserved.

I measured net N transformation rates of well-decayed boles and adjacent mineral soil under field conditions in an old-growth Douglas-fir/western hemlock/western red cedar stand in the central Oregon Cascades. Additionally, laboratory assays and in- cubations were used to elucidate the controls on net N transformations in these materials. Net N mineralization under field conditions was similar for well-decayed boles and mineral soils (0.83 and 0.61 g N m-2 material-yr-1, respectively). Laboratory rates of net N min- eralization per mass of total N were similar or higher in well-decayed boles compared to mineral soil. These results are surprising, given that the C:N ratio of well-decayed boles was much greater than that of mineral soil (117 vs. 26, respectively), and given the vastly different physical structure of these materials. Higher field and laboratory rates of net N mineralization relative to total N in boles compared to mineral soil suggest either that organically bound N contained in boles is in a more readily mineralizable form or that C compounds in well-decayed bole material are less readily metabolized by microbial het- erotrophs (i.e., C availability is lower) than in mineral soil, resulting in a higher ratio of net-to-gross N mineralization because of reduced N demand by a C-limited microflora. A lower microbial respiration rate and smaller microbial biomass C relative to the total C pool in well-decayed boles than in mineral soil support the latter hypothesis. Furthermore, bole material exhibited a higher specific respiration rate (respired C per unit microbial biomass C), suggesting either a lower C-use efficiency of bole microflora or a lower C availability in boles compared to mineral soil. Both well-decayed bole and mineral soil materials showed low annual rates of net nitrification under field conditions. Using an estimate of the mass of class 4 and 5 boles in similar forest stands in the Pacific Northwest, I estimate that well-decayed boles contribute about 0.16-0.25 g N.m-2 yr-I of plant-available N. This N flux is lower than other internal N fluxes within this forest, as well as lower than the rate of N input from the atmosphere. Total plant uptake in a nearby old-growth Douglas-fir forest has been estimated at about 4.0 g Nm-2_yr- 1, suggesting that well-decayed boles may contribute about 4-6% of plant N uptake. Results from this study indicate that the C:N ratio is a poor predictor of net N release from contrasting forest detrital pools.

We assessed the potential impact of global warming resulting from a doubling of preindustrial atmospheric CO2 on soil net N transformations by transferring intact soil cores (0–15 cm) from a high-elevation old-growth forest to a forest about 800 m lower in elevation in the central Oregon Cascade Mountains, USA. The lower elevation site had mean annual air and soil (10-cm mineral soil depth) temperatures about 2.4 and 3.9 °C higher than the high-elevation site, respectively. Annual rates of soil net N mineralization and nitrification more than doubled in soil transferred to the low-elevation site (17.2–36.0 kg N ha–1 and 5.0–10.7 kg NO3––N ha–1, respectively). Leaching of inorganic N from the surface soil (in the absence of plant uptake) also increased. The reciprocal treatment (transferring soil cores from the low- to the high-elevation site) resulted in decreases of about 70, 80, and 65% in annual rates of net N mineralization, nitrification, and inorganic N leaching, respectively. Laboratory incubations of soils under conditions of similar temperature and soil water potential suggest that the quality of soil organic matter is higher at the high-elevation site. Similar in situ rates of soil net N transformations between the two sites occurred because the lower temperature counteracts the effects of greater substrate quantity and quality at the high elevation site. Our results support the hypothesis that high-elevation, old-growth forest soils in the central Cascades have higher C and N storage than their low-elevation analogues primarily because low temperatures limit net C and N mineralization rates at higher elevations.

We measured surface soil (0-15cm) C and N pools and processes inside and outside an area that had been trenched 13 years earlier in an old-growth conifer forest (>450 years) to assess the long-term impacts of reduced root inputs on C and N turnover. Trenching, combined with frequent clipping of understory plants, was originally conducted to prevent nutrient uptake by plants, as part of a study of the role of vegetation in ecosystem retention of N. Thirteen years following trenching, the median values of bulk density, pH, total C and N concentrations, annual rates of in situ net N mineralization and nitrification, microbial biomass C and N, microbial respiration, and anaerobically mineralizable N in the trenched plot were all within the 25-75% interquartile range of values found in the replicated, untrenched plots. The trenched plot had higher rates of net N mineralization (41% higher in October, 484% higher in June) and net nitrification (25% higher in October, and lower net NO3- immobilization in June) during laboratory incubation and a 22% higher water content in October. In June, soil water content in the trenched plot was about 8% lower than in the untrenched plots. Our results suggest that soil C and N dynamics in these old-growth forests are relatively resistant to perturbations resulting from major reductions in root input to the soil.

In ponderosa pine (Pinus ponderosa Douglas ex P. Lawson & Lawson)-bunchgrass ecosystems of the western USA, fire exclusion by Euro-American settlers facilitated pine invasion of grassy openings, increased forest floor detritus, and shifted the disturbance regime toward stand-replacing fires, motivating ecological restoration through thinning and prescribed burning. We used in situ soil respiration over a 2-yr period to assess belowground responses to pine invasion and restoration in a ponderosa pine-bunchgrass ecosystem near Flagstaff, AZ. Replicated restoration treatments were: (i) partial restoration - thinning to presettlement conditions; (ii) complete restoration - removing trees and forest floor material to presettlement conditions, native grass litter addition, and prescribed burning; and (iii) control. Within treatments, we sampled beneath different canopy types to assess the effects of pine invasion into grassy openings on soil respiration. Growing season soil respiration was greater in the complete restoration (346 ± 24 g CO 2 -C m -2 ) and control (350 ± 8 g CO 2 -C m -2 ) than the partial restoration (301 ± 5 g CO 2 -C m -2 ) in 1995. In 1996, the complete (364 ± 17 g CO 2 -C m -2 ) and partial (328 ± 7 g CO 2 -C m -2 ) restoration treatments had greater growing season respiration rates than the control (302 ± 13 g CO 2 -C m -2 ). Results suggest that restoration effects on soil respiration depend on interannual soil water patterns and may not significantly alter regional C cydes. Soil respiration from grassy openings was 15% greater than from soil beneath presettlement or postsettlement pines in 1995 and 1996. A lack of active management will decrease belowground catabolism if pines continue to expand at the expense of grassy openings.

THE importance of nitrate (NO–3) in the internal nitrogen cycle of undisturbed coniferous ecosystems has not been widely recognized1,2. Nitrate concentrations in soils from these forests tend to be low, and assays measuring net nitrification usually show exceedingly slow rates3,4. It may be, however, that microbial assimilation of NO–3 is substantial in these soils, and that net nitrification rates greatly underestimate gross rates5. Here we use a 15N isotope-dilution technique in intact soil cores to measure gross rates of nitrification and microbial assimilation of NO–3 in eleven undisturbed forest ecosystems of New Mexico and Oregon. We found that gross nitrification rates were surprisingly high in all of the forests examined. Net nitrification rates poorly predicted gross rates because the soil microbial communities had the capacity to assimilate almost all of the NO–3 produced. To our knowledge, this is the first report of gross nitrification and NO–3 assimilation rates in intact soil samples from a large number of contrasting forest ecosystems. Our results contradict previous assumptions that nitrification rates are low in mature coniferous forests and suggest that current models greatly underestimate the role of the microbial community in preventing NO–3 loss.

We present a significant relationship between the natural abundance isotopic composition of ecosystem pools and the abundance of a microbial gene. Natural abundance 15N of soils and soil DNA were analysed and compared with archaeal ammonia oxidizer abundance along an elevation gradient in northern Arizona and along a substrate age gradient in Hawai'i. There was a significant positive correlation between the abundance of archaeal amoA genes and natural abundance δ15N of total soil or DNA suggesting that ammonia oxidizing archaea play an important role in ecosystem N release. © 2013 Elsevier Ltd.

A comparative study of N dynamics in an annual grassland and a young mixed-conifer forest, that occur on the same soil great group in California, was made by adding small amounts of 15NH4+ to mineral soils within field microplots and following changes in 15N recoveries in soil and plant pools over 12–16 months. In annual grassland microplots, the aboveground plant biomass contained 5–6% of the added 15N at the end of the growing season. One month after 15N addition almost 70% of the added 15N was recovered in the soil organic matter (SOM) pool, which included surface litter and roots but not microbial biomass. Laboratory incubation of sterilized soil cores indicated that as much as half of the 15N recovered in the SOM pool may have been fixed abiotically. Nevertheless, substantial and rapid incorporation of 15N in the SOM pool indicates a high potential for N immobilization and a rapid turnover of the microbial biomass-N pool in this soil. Recovery of 15N in microbial biomass ranged from 9 to 15%; recovery in the soil solution was lower than in the microbial biomass, but showed a similar seasonal trend. Total 15N recovery ranged from 72 to 85% over the study period. In forest microplots (which excluded plants), recovery of added 15N was generally lower than in the grassland for all soil pools examined. Recovery of 15N in SOM and microbial pools showed reciprocal trends seasonally, suggesting that there was considerable movement of N between these pools in the forest soil. Total 15N recovery was only about 54% 6 months after 15N addition, indicating a high potential for rapid N loss from the surface soil of this forest. Substantially lower recovery of 15N in the forest soil cannot be attributed entirely to the exclusion of plant uptake because plant 15N recovery in the grassland was relatively low. The low 15N recovery in the forest soil might be due to the low capacity of this soil to immobilize N and protect immobilized N from being remineralized and lost from the soil via leaching or denitrification. In both ecosystems estimates of N flow from the mineral soil to decomposing surface residues (presumably via fungal translocation) was significant (0.29 and 0.02 g-N m−2 yr−1 for the grassland and forest, respectively) relative to the net internal dynamics of N within the surface detritus. Comparison of net changes in litter N and accumulation of 15N during decomposition suggests that N was mineralized from litter concurrently as N was immobilized from the mineral soil in both sites.

Better understanding of variation in soil carbon dioxide (CO2) efflux caused by measurement techniques is needed, especially over gradients of site disturbance, to accurately estimate the global carbon cycle. We present soil CO2 efflux data from a gradient of disturbance to ponderosa pine (Pinus ponderosa C. Lawson var. scopulorum Engelm.) forests in northern Arizona, USA that were obtained using four different techniques: vented static chambers, a Licor 6400-09, and soil CO2 diffusion profiles using two different models (Moldrup, Millington–Quirk) to estimate soil gas diffusivity. We also compared soil CO2 efflux measured by the Moldrup and Millington–Quirk diffusion profile methods to nighttime total ecosystem respiration (TER) data from an eddy covariance tower. We addressed four questions: (1) Does the use of a given method to measure soil CO2 efflux bias results across a disturbance gradient? (2) Does the magnitude of difference between observed and modeled estimates of soil CO2 differ between methods and across sites? (3) What is the spatial variability of each method at each site? (4) Which method is closest to the estimate of TER measured by the eddy covariance tower? Although soil CO2 efflux varied significantly among methods the differences were consistent among sites. Measured and modeled total growing season fluxes were generally higher for the Licor 6400-09 and Millington–Quirk diffusion gradient methods compared with static chamber and the Moldrup diffusion gradient methods. A power analysis showed that the larger static chamber was the most efficient method at sampling spatial variation in soil CO2 efflux. Nighttime measurements of soil CO2 efflux from the Moldrup diffusion gradient method were most strongly related to nighttime TER assessed with eddy covariance. The use of a single, well-implemented method to measure soil CO2 efflux is unlikely to create bias in comparisons across a gradient of forest disturbance.