Urbanization and edge effects interact to drive mutualism breakdown and the rise of unstable pathogenic communities in forest soil.

Temperate forests are threatened by urbanization and fragmentation, with over 20% (118,300 km2) of U.S. forest land projected to be subsumed by urban land development. We leveraged a unique, well-characterized urban-to-rural and forest edge-to-interior gradient to identify the combined impact of these two land use changes-urbanization and forest edge creation-on the soil microbial community in native remnant forests. We found evidence of mutualism breakdown between trees and their fungal root mutualists [ectomycorrhizal (ECM) fungi] with urbanization, where ECM fungi colonized fewer tree roots and had less connectivity in soil microbiome networks in urban forests compared to rural forests. However, urbanization did not reduce the relative abundance of ECM fungi in forest soils; instead, forest edges alone led to strong reductions in ECM fungal abundance. At forest edges, ECM fungi were replaced by plant and animal pathogens, as well as copiotrophic, xenobiotic-degrading, and nitrogen-cycling bacteria, including nitrifiers and denitrifiers. Urbanization and forest edges interacted to generate new "suites" of microbes, with urban interior forests harboring highly homogenized microbiomes, while edge forest microbiomes were more heterogeneous and less stable, showing increased vulnerability to low soil moisture. When scaled to the regional level, we found that forest soils are projected to harbor high abundances of fungal pathogens and denitrifying bacteria, even in rural areas, due to the widespread existence of forest edges. Our results highlight the potential for soil microbiome dysfunction-including increased greenhouse gas production-in temperate forest regions that are subsumed by urban expansion, both now and in the future.

Soils at the temperate forest edge: An investigation of soil characteristics and carbon dynamics.

Global proliferation of forest edges through anthropogenic land-use change and forest fragmentation is well documented, and while forest fragmentation has clear consequences for soil carbon (C) cycling, underlying drivers of belowground activity at the forest edge remain poorly understood. Increasing soil C losses via respiration have been observed at rural forest edges, but this process was suppressed at urban forest edges. We offer a comprehensive, coupled investigation of abiotic soil conditions and biotic soil activity from forest edge to interior at eight sites along an urbanization gradient to elucidate how environmental stressors are linked to soil C cycling at the forest edge. Despite significant diverging trends in edge soil C losses between urban and rural sites, we did not find comparable differences in soil % C or microbial enzyme activity, suggesting an unexpected decoupling of soil C fluxes and pools at forest edges. We demonstrate that across site types, soils at forest edges were less acidic than the forest interior (p < 0.0001), and soil pH was positively correlated with soil calcium, magnesium and sodium content (adj R2 = 0.37), which were also elevated at the edge. Compared to forest interior, forest edge soils exhibited a 17.8 % increase in sand content and elevated freeze-thaw frequency with probable downstream effects on root turnover and decomposition. Using these and other novel forest edge data, we demonstrate that significant variation in edge soil respiration (adj R2 = 0.46; p = 0.0002) and C content (adj R2 = 0.86; p < 0.0001) can be explained using soil parameters often mediated by human activity (e.g., soil pH, trace metal and cation concentrations, soil temperature), and we emphasize the complex influence of multiple, simultaneous global change drivers at forest edges. Forest edge soils reflect legacies of anthropogenic land-use and modern human management, and this must be accounted for to understand soil activity and C cycling across fragmented landscapes.

Applied winter biology: threats, conservation and management of biological resources during winter in cold climate regions.

Winter at high latitudes is characterized by low temperatures, dampened light levels and short photoperiods which shape ecological and evolutionary outcomes from cells to populations to ecosystems. Advances in our understanding of winter biological processes (spanning physiology, behaviour and ecology) highlight that biodiversity threats (e.g. climate change driven shifts in reproductive windows) may interact with winter conditions, leading to greater ecological impacts. As such, conservation and management strategies that consider winter processes and their consequences on biological mechanisms may lead to greater resilience of high altitude and latitude ecosystems. Here, we use well-established threat and action taxonomies produced by the International Union of Conservation of Nature-Conservation Measures Partnership (IUCN-CMP) to synthesize current threats to biota that emerge during, or as the result of, winter processes then discuss targeted management approaches for winter-based conservation. We demonstrate the importance of considering winter when identifying threats to biodiversity and deciding on appropriate management strategies across species and ecosystems. We confirm our expectation that threats are prevalent during the winter and are especially important considering the physiologically challenging conditions that winter presents. Moreover, our findings emphasize that climate change and winter-related constraints on organisms will intersect with other stressors to potentially magnify threats and further complicate management. Though conservation and management practices are less commonly considered during the winter season, we identified several potential or already realized applications relevant to winter that could be beneficial. Many of the examples are quite recent, suggesting a potential turning point for applied winter biology. This growing body of literature is promising but we submit that more research is needed to identify and address threats to wintering biota for targeted and proactive conservation. We suggest that management decisions consider the importance of winter and incorporate winter specific strategies for holistic and mechanistic conservation and resource management.

Urbanization and fragmentation have opposing effects on soil nitrogen availability in temperate forest ecosystems.

Nitrogen (N) availability relative to plant demand has been declining in recent years in terrestrial ecosystems throughout the world, a phenomenon known as N oligotrophication. The temperate forests of the northeastern U.S. have experienced a particularly steep decline in bioavailable N, which is expected to be exacerbated by climate change. This region has also experienced rapid urban expansion in recent decades that leads to forest fragmentation, and it is unknown whether and how these changes affect N availability and uptake by forest trees. Many studies have examined the impact of either urbanization or forest fragmentation on nitrogen (N) cycling, but none to our knowledge have focused on the combined effects of these co-occurring environmental changes. We examined the effects of urbanization and fragmentation on oak-dominated (Quercus spp.) forests along an urban to rural gradient from Boston to central Massachusetts (MA). At eight study sites along the urbanization gradient, plant and soil measurements were made along a 90 m transect from a developed edge to an intact forest interior. Rates of net ammonification, net mineralization, and foliar N concentrations were significantly higher in urban than rural sites, while net nitrification and foliar C:N were not different between urban and rural forests. At urban sites, foliar N and net ammonification and mineralization were higher at forest interiors compared to edges, while net nitrification and foliar C:N were higher at rural forest edges than interiors. These results indicate that urban forests in the northeastern U.S. have greater soil N availability and N uptake by trees compared to rural forests, counteracting the trend for widespread N oligotrophication in temperate forests around the globe. Such increases in available N are diminished at forest edges, however, demonstrating that forest fragmentation has the opposite effect of urbanization on coupled N availability and demand by trees.

Evidence, causes, and consequences of declining nitrogen availability in terrestrial ecosystems.

The productivity of ecosystems and their capacity to support life depends on access to reactive nitrogen (N). Over the past century, humans have more than doubled the global supply of reactive N through industrial and agricultural activities. However, long-term records demonstrate that N availability is declining in many regions of the world. Reactive N inputs are not evenly distributed, and global changes-including elevated atmospheric carbon dioxide (CO2) levels and rising temperatures-are affecting ecosystem N supply relative to demand. Declining N availability is constraining primary productivity, contributing to lower leaf N concentrations, and reducing the quality of herbivore diets in many ecosystems. We outline the current state of knowledge about declining N availability and propose actions aimed at characterizing and responding to this emerging challenge.

Diverging patterns at the forest edge: Soil respiration dynamics of fragmented forests in urban and rural areas.

As urbanization and forest fragmentation increase around the globe, it is critical to understand how rates of respiration and carbon losses from soil carbon pools are affected by these processes. This study characterizes soils in fragmented forests along an urban to rural gradient, evaluating the sensitivity of soil respiration to changes in soil temperature and moisture near the forest edge. While previous studies found elevated rates of soil respiration at temperate forest edges in rural areas compared to the forest interior, we find that soil respiration is suppressed at the forest edge in urban areas. At urban sites, respiration rates are 25% lower at the forest edge relative to the interior, likely due to high temperature and aridity conditions near urban edges. While rural soils continue to respire with increasing temperatures, urban soil respiration rates asymptote as temperatures climb and soils dry. Soil temperature- and moisture-sensitivity modeling shows that respiration rates in urban soils are less sensitive to rising temperatures than those in rural soils. Scaling these results to Massachusetts (MA), which encompasses 0.25 Mha of the urban forest, we find that failure to account for decreases in soil respiration rates near urban forest edges leads to an overestimate of growing-season soil carbon fluxes of >350,000 Mg C. This difference is almost 2.5 times that for rural soils in the analogous comparison (underestimate of <143,000 Mg C), even though rural forest area is more than four times greater than urban forest area in MA. While a changing climate may stimulate carbon losses from rural forest edge soils, urban forests may experience enhanced soil carbon sequestration near the forest edge. These findings highlight the need to capture the effects of forest fragmentation and land use context when making projections about soil behavior and carbon cycling in a warming and increasingly urbanized world.

Influence of landscape management practices on urban greenhouse gas budgets.

BACKGROUND: With a lack of United States federal policy to address climate change, cities, the private sector, and universities have shouldered much of the work to reduce carbon dioxide (CO2) and other greenhouse gas emissions. This study aims to determine how landcover characteristics influence the amount of carbon (C) sequestered and respired via biological processes, evaluating the role of land management on the overall C budget of an urban university. Boston University published a comprehensive Climate Action Plan in 2017 with the goal of achieving C neutrality by 2040. In this study, we digitized and discretized each of Boston University's three urban campuses into landcover types, with C sequestration and respiration rates measured and scaled to provide a University-wide estimate of biogenic C fluxes within the broader context of total University emissions. RESULTS: Each of Boston University's three highly urban campuses were net sources of biogenic C to the atmosphere. While trees were estimated to sequester 0.6 ± 0.2 kg C m-2 canopy cover year-1, mulch and lawn areas in 2018 emitted C at rates of 1.7 ± 0.4 kg C m-2 year-1 and 1.4 ± 0.4 kg C m-2 year-1, respectively. C uptake by tree canopy cover, which can spatially overlap lawn and mulched landcovers, was not large enough to offset biogenic emissions. The proportion of biogenic emissions to Scope 1 anthropogenic emissions on each campus varied from 0.5% to 2%, and depended primarily on the total anthropogenic emissions on each campus. CONCLUSIONS: Our study quantifies the role of urban landcover in local C budgets, offering insights on how landscaping management strategies-such as decreasing mulch application rates and expanding tree canopy extent-can assist universities in minimizing biogenic C emissions and even potentially creating a small biogenic C sink. Although biogenic C fluxes represent a small fraction of overall anthropogenic emissions on urban university campuses, these biogenic fluxes are under active management by the university and should be included in climate action plans.

Growing-season warming and winter soil freeze/thaw cycles increase transpiration in a northern hardwood forest.

Climate models project higher growing-season temperatures and a decline in the depth and duration of winter snowpack throughout many north temperate ecosystems over the next century. A smaller snowpack is projected to induce more frequent soil freeze/thaw cycles in winter in northern hardwood forests of the northeastern United States. We measured the combined effects of warmer growing-season soil temperatures and increased winter freeze/thaw cycles on rates of leaf-level photosynthesis and transpiration (sap flow) of red maple (Acer rubrum) trees in a northern hardwood forest at the Climate Change Across Seasons Experiment at Hubbard Brook Experimental Forest in New Hampshire. Soil temperatures were warmed 5°C above ambient temperatures during the growing season and soil freeze/thaw cycles were induced in winter to mimic the projected changes in soil temperature over the next century. Relative to reference plots, growing-season soil warming increased rates of leaf-level photosynthesis by up to 85.32 ± 4.33%, but these gains were completely offset by soil freeze/thaw cycles in winter, suggesting that increased freeze/thaw cycles in winter over the next 100 yr will reduce the effect of warming on leaf-level carbon gains. Soil warming in the growing season increased rates of transpiration per kilopascal of vapor pressure deficit (VPD) by up to 727.39 ± 0.28%, even when trees were exposed to increased frequency of soil freeze/thaw cycles in the previous winter, which could influence regional hydrology in the future. Using climate projections downscaled from the Coupled Model Intercomparison Project, we project increased rates of whole-season transpiration in these forests over the next century by 42-61%. We also project 52-77 additional days when daily air temperatures will be above the long-term average daily maximum during the growing season at Hubbard Brook. Together, these results show that projected changes in climate across both the growing season and winter are likely to cause greater rates of water uptake and have no effect on rates of leaf-level carbon uptake by trees, with potential ecosystem consequences for hydrology and carbon cycling in northern hardwood forests.

Soil Microbes Trade-Off Biogeochemical Cycling for Stress Tolerance Traits in Response to Year-Round Climate Change.

Winter air temperatures are rising faster than summer air temperatures in high-latitude forests, increasing the frequency of soil freeze/thaw events in winter. To determine how climate warming and soil freeze/thaw cycles affect soil microbial communities and the ecosystem processes they drive, we leveraged the Climate Change across Seasons Experiment (CCASE) at the Hubbard Brook Experimental Forest in the northeastern United States, where replicate field plots receive one of three climate treatments: warming (+5°C above ambient in the growing season), warming in the growing season + winter freeze/thaw cycles (+5°C above ambient +4 freeze/thaw cycles during winter), and no treatment. Soil samples were taken from plots at six time points throughout the growing season and subjected to amplicon (rDNA) and metagenome sequencing. We found that soil fungal and bacterial community composition were affected by changes in soil temperature, where the taxonomic composition of microbial communities shifted more with the combination of growing-season warming and increased frequency of soil freeze/thaw cycles in winter than with warming alone. Warming increased the relative abundance of brown rot fungi and plant pathogens but decreased that of arbuscular mycorrhizal fungi, all of which recovered under combined growing-season warming and soil freeze/thaw cycles in winter. The abundance of animal parasites increased significantly under combined warming and freeze/thaw cycles. We also found that warming and soil freeze/thaw cycles suppressed bacterial taxa with the genetic potential for carbon (i.e., cellulose) decomposition and soil nitrogen cycling, such as N fixation and the final steps of denitrification. These new soil communities had higher genetic capacity for stress tolerance and lower genetic capacity to grow or reproduce, relative to the communities exposed to warming in the growing season alone. Our observations suggest that initial suppression of biogeochemical cycling with year-round climate change may be linked to the emergence of taxa that trade-off growth for stress tolerance traits.

A meta-analysis of 1,119 manipulative experiments on terrestrial carbon-cycling responses to global change.

Direct quantification of terrestrial biosphere responses to global change is crucial for projections of future climate change in Earth system models. Here, we synthesized ecosystem carbon-cycling data from 1,119 experiments performed over the past four decades concerning changes in temperature, precipitation, CO2 and nitrogen across major terrestrial vegetation types of the world. Most experiments manipulated single rather than multiple global change drivers in temperate ecosystems of the USA, Europe and China. The magnitudes of warming and elevated CO2 treatments were consistent with the ranges of future projections, whereas those of precipitation changes and nitrogen inputs often exceeded the projected ranges. Increases in global change drivers consistently accelerated, but decreased precipitation slowed down carbon-cycle processes. Nonlinear (including synergistic and antagonistic) effects among global change drivers were rare. Belowground carbon allocation responded negatively to increased precipitation and nitrogen addition and positively to decreased precipitation and elevated CO2. The sensitivities of carbon variables to multiple global change drivers depended on the background climate and ecosystem condition, suggesting that Earth system models should be evaluated using site-specific conditions for best uses of this large dataset. Together, this synthesis underscores an urgent need to explore the interactions among multiple global change drivers in underrepresented regions such as semi-arid ecosystems, forests in the tropics and subtropics, and Arctic tundra when forecasting future terrestrial carbon-climate feedback.

Roots Mediate the Effects of Snowpack Decline on Soil Bacteria, Fungi, and Nitrogen Cycling in a Northern Hardwood Forest.

Rising winter air temperature will reduce snow depth and duration over the next century in northern hardwood forests. Reductions in snow depth may affect soil bacteria and fungi directly, but also affect soil microbes indirectly through effects of snowpack loss on plant roots. We incubated root exclusion and root ingrowth cores across a winter climate-elevation gradient in a northern hardwood forest for 29 months to identify direct (i.e., winter snow-mediated) and indirect (i.e., root-mediated) effects of winter snowpack decline on soil bacterial and fungal communities, as well as on potential nitrification and net N mineralization rates. Both winter snowpack decline and root exclusion increased bacterial richness and phylogenetic diversity. Variation in bacterial community composition was best explained by differences in winter snow depth or soil frost across elevation. Root ingrowth had a positive effect on the relative abundance of several bacterial taxonomic orders (e.g., Acidobacterales and Actinomycetales). Nominally saprotrophic (e.g., Saccharomycetales and Mucorales) or mycorrhizal (e.g., Helotiales, Russalales, Thelephorales) fungal taxonomic orders were also affected by both root ingrowth and snow depth variation. However, when grouped together, the relative abundance of saprotrophic fungi, arbuscular mycorrhizal fungi, and ectomycorrhizal fungi were not affected by root ingrowth or snow depth, suggesting that traits in addition to trophic mode will mediate fungal community responses to snowpack decline in northern hardwood forests. Potential soil nitrification rates were positively related to ammonia-oxidizing bacteria and archaea abundance (e.g., Nitrospirales, Nitrosomondales, Nitrosphaerales). Rates of N mineralization were positively and negatively correlated with ectomycorrhizal and saprotrophic fungi, respectively, and these relationships were mediated by root exclusion. The results from this study suggest that a declining winter snowpack and its effect on plant roots each have direct effects on the diversity and abundance of soil bacteria and fungal communities that interact to determine rates of soil N cycling in northern hardwood forests.

Declines in northern forest tree growth following snowpack decline and soil freezing.

Changes in growing season climate are often the foci of research exploring forest response to climate change. By contrast, little is known about tree growth response to projected declines in winter snowpack and increases in soil freezing in seasonally snow-covered forest ecosystems, despite extensive documentation of the importance of winter climate in mediating ecological processes. We conducted a 5-year snow-removal experiment whereby snow was removed for the first 4-5 weeks of winter in a northern hardwood forest at the Hubbard Brook Experimental Forest in New Hampshire, USA. Our results indicate that adverse impacts of reduced snowpack and increased soil freezing on the physiology of Acer saccharum (sugar maple), a dominant species across northern temperate forests, are accompanied by a 40 ± 3% reduction in aboveground woody biomass increment, averaged across the 6 years following the start of the experiment. Further, we find no indication of growth recovery 1 year after cessation of the experiment. Based on these findings, we integrate spatial modeling of snowpack depth with forest inventory data to develop a spatially explicit, regional-scale assessment of the vulnerability of forest aboveground growth to projected declines in snowpack depth and increased soil frost. These analyses indicate that nearly 65% of sugar maple basal area in the northeastern United States resides in areas that typically experience insulating snowpack. However, under the RCP 4.5 and 8.5 emissions scenarios, we project a 49%-95% reduction in forest area experiencing insulating snowpack by the year 2099 in the northeastern United States, leaving large areas of northern forest vulnerable to these changes in winter climate, particularly along the northern edge of the region. Our study demonstrates that research focusing on growing season climate alone overestimates the stimulatory effect of warming temperatures on tree and forest growth in seasonally snow-covered forests.

Isotopic evidence for oligotrophication of terrestrial ecosystems.

Human societies depend on an Earth system that operates within a constrained range of nutrient availability, yet the recent trajectory of terrestrial nitrogen (N) availability is uncertain. Examining patterns of foliar N concentrations and isotope ratios (δ15N) from more than 43,000 samples acquired over 37 years, here we show that foliar N concentration declined by 9% and foliar δ15N declined by 0.6-1.6‰. Examining patterns across different climate spaces, foliar δ15N declined across the entire range of mean annual temperature and mean annual precipitation tested. These results suggest declines in N supply relative to plant demand at the global scale. In all, there are now multiple lines of evidence of declining N availability in many unfertilized terrestrial ecosystems, including declines in δ15N of tree rings and leaves from herbarium samples over the past 75-150 years. These patterns are consistent with the proposed consequences of elevated atmospheric carbon dioxide and longer growing seasons. These declines will limit future terrestrial carbon uptake and increase nutritional stress for herbivores.

What Goes Up Must Come Down: Integrating Air and Water Quality Monitoring for Nutrients.

Excess nitrogen and phosphorus ("nutrients") loadings continue to affect ecosystem function and human health across the U.S. Our ability to connect atmospheric inputs of nutrients to aquatic end points remains limited due to uncoupled air and water quality monitoring. Where connections exist, the information provides insights about source apportionment, trends, risk to sensitive ecosystems, and efficacy of pollution reduction efforts. We examine several issues driving the need for better integrated monitoring, including: coastal eutrophication, urban hotspots of deposition, a shift from oxidized to reduced nitrogen deposition, and the disappearance of pristine lakes. Successful coordination requires consistent data reporting; collocating deposition and water quality monitoring; improving phosphorus deposition measurements; and filling coverage gaps in urban corridors, agricultural areas, undeveloped watersheds, and coastal zones.

Variability, drivers, and effects of atmospheric nitrogen inputs across an urban area: Emerging patterns among human activities, the atmosphere, and soils.

Atmospheric deposition of nitrogen (N) is a major input of N to the biosphere and is elevated beyond preindustrial levels throughout many ecosystems. Deposition monitoring networks in the United States generally avoid urban areas in order to capture regional patterns of N deposition, and studies measuring N deposition in cities usually include only one or two urban sites in an urban-rural comparison or as an anchor along an urban-to-rural gradient. Describing patterns and drivers of atmospheric N inputs is crucial for understanding the effects of N deposition; however, little is known about the variability and drivers of atmospheric N inputs or their effects on soil biogeochemistry within urban ecosystems. We measured rates of canopy throughfall N as a measure of atmospheric N inputs, as well as soil net N mineralization and nitrification, soil solution N, and soil respiration at 15 sites across the greater Boston, Massachusetts area. Rates of throughfall N are 8.70±0.68kgNha-1yr-1, vary 3.5-fold across sites, and are positively correlated with rates of local vehicle N emissions. Ammonium (NH4+) composes 69.9±2.2% of inorganic throughfall N inputs and is highest in late spring, suggesting a contribution from local fertilizer inputs. Soil solution NO3- is positively correlated with throughfall NO3- inputs. In contrast, soil solution NH4+, net N mineralization, nitrification, and soil respiration are not correlated with rates of throughfall N inputs. Rather, these processes are correlated with soil properties such as soil organic matter. Our results demonstrate high variability in rates of urban throughfall N inputs, correlation of throughfall N inputs with local vehicle N emissions, and a decoupling of urban soil biogeochemistry and throughfall N inputs.

Climate Change Across Seasons Experiment (CCASE): A new method for simulating future climate in seasonally snow-covered ecosystems.

Climate models project an increase in mean annual air temperatures and a reduction in the depth and duration of winter snowpack for many mid and high latitude and high elevation seasonally snow-covered ecosystems over the next century. The combined effects of these changes in climate will lead to warmer soils in the growing season and increased frequency of soil freeze-thaw cycles (FTCs) in winter due to the loss of a continuous, insulating snowpack. Previous experiments have warmed soils or removed snow via shoveling or with shelters to mimic projected declines in the winter snowpack. To our knowledge, no experiment has examined the interactive effects of declining snowpack and increased frequency of soil FTCs, combined with soil warming in the snow-free season on terrestrial ecosystems. In addition, none have mimicked directly the projected increase in soil FTC frequency in tall statured forests that is expected as a result of a loss of insulating snow in winter. We established the Climate Change Across Seasons Experiment (CCASE) at Hubbard Brook Experimental Forest in the White Mountains of New Hampshire in 2012 to assess the combined effects of these changes in climate on a variety of pedoclimate conditions, biogeochemical processes, and ecology of northern hardwood forests. This paper demonstrates the feasibility of creating soil FTC events in a tall statured ecosystem in winter to simulate the projected increase in soil FTC frequency over the next century and combines this projected change in winter climate with ecosystem warming throughout the snow-free season. Together, this experiment provides a new and more comprehensive approach for climate change experiments that can be adopted in other seasonally snow-covered ecosystems to simulate expected changes resulting from global air temperature rise.

Ecosystem fluxes of hydrogen in a mid-latitude forest driven by soil microorganisms and plants.

Molecular hydrogen (H2 ) is an atmospheric trace gas with a large microbe-mediated soil sink, yet cycling of this compound throughout ecosystems is poorly understood. Measurements of the sources and sinks of H2 in various ecosystems are sparse, resulting in large uncertainties in the global H2 budget. Constraining the H2 cycle is critical to understanding its role in atmospheric chemistry and climate. We measured H2 fluxes at high frequency in a temperate mixed deciduous forest for 15 months using a tower-based flux-gradient approach to determine both the soil-atmosphere and the net ecosystem flux of H2 . We found that Harvard Forest is a net H2 sink (-1.4 ± 1.1 kg H2  ha-1 ) with soils as the dominant H2 sink (-2.0 ± 1.0 kg H2  ha-1 ) and aboveground canopy emissions as the dominant H2 source (+0.6 ± 0.8 kg H2  ha-1 ). Aboveground emissions of H2 were an unexpected and substantial component of the ecosystem H2 flux, reducing net ecosystem uptake by 30% of that calculated from soil uptake alone. Soil uptake was highly seasonal (July maximum, February minimum), positively correlated with soil temperature and negatively correlated with environmental variables relevant to diffusion into soils (i.e., soil moisture, snow depth, snow density). Soil microbial H2 uptake was correlated with rhizosphere respiration rates (r = 0.8, P < 0.001), and H2 metabolism yielded up to 2% of the energy gleaned by microbes from carbon substrate respiration. Here, we elucidate key processes controlling the biosphere-atmosphere exchange of H2 and raise new questions regarding the role of aboveground biomass as a source of atmospheric H2 and mechanisms linking soil H2 and carbon cycling. Results from this study should be incorporated into modeling efforts to predict the response of the H2 soil sink to changes in anthropogenic H2 emissions and shifting soil conditions with climate and land-use change.

Reduced snow cover alters root-microbe interactions and decreases nitrification rates in a northern hardwood forest.

Snow cover is projected to decline during the next century in many ecosystems that currently experience a seasonal snowpack. Because snow insulates soils from frigid winter air temperatures, soils are expected to become colder and experience more winter soil freeze-thaw cycles as snow cover continues to decline. Tree roots are adversely affected by snowpack reduction, but whether loss of snow will affect root-microbe interactions remains largely unknown. The objective of this study was to distinguish and attribute direct (e.g., winter snow- and/or soil frost-mediated) vs. indirect (e.g., root-mediated) effects of winter climate change on microbial biomass, the potential activity of microbial exoenzymes, and net N mineralization and nitrification rates. Soil cores were incubated in situ in nylon mesh that either allowed roots to grow into the soil core (2 mm pore size) or excluded root ingrowth (50 µm pore size) for up to 29 months along a natural winter climate gradient at Hubbard Brook Experimental Forest, NH (USA). Microbial biomass did not differ among ingrowth or exclusion cores. Across sampling dates, the potential activities of cellobiohydrolase, phenol oxidase, and peroxidase, and net N mineralization rates were more strongly related to soil volumetric water content (P < 0.05; R2  = 0.25-0.46) than to root biomass, snow or soil frost, or winter soil temperature (R2  < 0.10). Root ingrowth was positively related to soil frost (P < 0.01; R2  = 0.28), suggesting that trees compensate for overwinter root mortality caused by soil freezing by re-allocating resources towards root production. At the sites with the deepest snow cover, root ingrowth reduced nitrification rates by 30% (P < 0.01), showing that tree roots exert significant influence over nitrification, which declines with reduced snow cover. If soil freezing intensifies over time, then greater compensatory root growth may reduce nitrification rates directly via plant-microbe N competition and indirectly through a negative feedback on soil moisture, resulting in lower N availability to trees in northern hardwood forests.