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.

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.

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.

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.

Temperature response of soil respiration largely unaltered with experimental warming.

The respiratory release of carbon dioxide (CO2) from soil is a major yet poorly understood flux in the global carbon cycle. Climatic warming is hypothesized to increase rates of soil respiration, potentially fueling further increases in global temperatures. However, despite considerable scientific attention in recent decades, the overall response of soil respiration to anticipated climatic warming remains unclear. We synthesize the largest global dataset to date of soil respiration, moisture, and temperature measurements, totaling >3,800 observations representing 27 temperature manipulation studies, spanning nine biomes and over 2 decades of warming. Our analysis reveals no significant differences in the temperature sensitivity of soil respiration between control and warmed plots in all biomes, with the exception of deserts and boreal forests. Thus, our data provide limited evidence of acclimation of soil respiration to experimental warming in several major biome types, contrary to the results from multiple single-site studies. Moreover, across all nondesert biomes, respiration rates with and without experimental warming follow a Gaussian response, increasing with soil temperature up to a threshold of ∼25 °C, above which respiration rates decrease with further increases in temperature. This consistent decrease in temperature sensitivity at higher temperatures demonstrates that rising global temperatures may result in regionally variable responses in soil respiration, with colder climates being considerably more responsive to increased ambient temperatures compared with warmer regions. Our analysis adds a unique cross-biome perspective on the temperature response of soil respiration, information critical to improving our mechanistic understanding of how soil carbon dynamics change with climatic warming.

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.

Seasonal fluxes of carbonyl sulfide in a midlatitude forest.

Carbonyl sulfide (OCS), the most abundant sulfur gas in the atmosphere, has a summer minimum associated with uptake by vegetation and soils, closely correlated with CO2. We report the first direct measurements to our knowledge of the ecosystem flux of OCS throughout an annual cycle, at a mixed temperate forest. The forest took up OCS during most of the growing season with an overall uptake of 1.36 ± 0.01 mol OCS per ha (43.5 ± 0.5 g S per ha, 95% confidence intervals) for the year. Daytime fluxes accounted for 72% of total uptake. Both soils and incompletely closed stomata in the canopy contributed to nighttime fluxes. Unexpected net OCS emission occurred during the warmest weeks in summer. Many requirements necessary to use fluxes of OCS as a simple estimate of photosynthesis were not met because OCS fluxes did not have a constant relationship with photosynthesis throughout an entire day or over the entire year. However, OCS fluxes provide a direct measure of ecosystem-scale stomatal conductance and mesophyll function, without relying on measures of soil evaporation or leaf temperature, and reveal previously unseen heterogeneity of forest canopy processes. Observations of OCS flux provide powerful, independent means to test and refine land surface and carbon cycle models at the ecosystem scale.

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.

Global patterns and substrate-based mechanisms of the terrestrial nitrogen cycle.

Nitrogen (N) deposition is impacting the services that ecosystems provide to humanity. However, the mechanisms determining impacts on the N cycle are not fully understood. To explore the mechanistic underpinnings of N impacts on N cycle processes, we reviewed and synthesised recent progress in ecosystem N research through empirical studies, conceptual analysis and model simulations. Experimental and observational studies have revealed that the stimulation of plant N uptake and soil retention generally diminishes as N loading increases, while dissolved and gaseous losses of N occur at low N availability but increase exponentially and become the dominant fate of N at high loading rates. The original N saturation hypothesis emphasises sequential N saturation from plant uptake to soil retention before N losses occur. However, biogeochemical models that simulate simultaneous competition for soil N substrates by multiple processes match the observed patterns of N losses better than models based on sequential competition. To enable better prediction of terrestrial N cycle responses to N loading, we recommend that future research identifies the response functions of different N processes to substrate availability using manipulative experiments, and incorporates the measured N saturation response functions into conceptual, theoretical and quantitative analyses.

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.

The promise and peril of intensive-site-based ecological research: insights from the Hubbard Brook ecosystem study.

Ecological research is increasingly concentrated at particular locations or sites. This trend reflects a variety of advantages of intensive, site-based research, but also raises important questions about the nature of such spatially delimited research: how well does site based research represent broader areas, and does it constrain scientific discovery? We provide an overview of these issues with a particular focus on one prominent intensive research site: the Hubbard Brook Experimental Forest (HBEF), New Hampshire, USA. Among the key features of intensive sites are: long-term, archived data sets that provide a context for new discoveries and the elucidation of ecological mechanisms; the capacity to constrain inputs and parameters, and to validate models of complex ecological processes; and the intellectual cross-fertilization among disciplines in ecological and environmental sciences. The feasibility of scaling up ecological observations from intensive sites depends upon both the phenomenon of interest and the characteristics of the site. An evaluation of deviation metrics for the HBEF illustrates that, in some respects, including sensitivity and recovery of streams and trees from acid deposition, this site is representative of the Northern Forest region, of which HBEF is a part. However, the mountainous terrain and lack of significant agricultural legacy make the HBEF among the least disturbed sites in the Northern Forest region. Its relatively cool, wet climate contributes to high stream flow compared to other sites. These similarities and differences between the HBEF and the region can profoundly influence ecological patterns and processes and potentially limit the generality of observations at this and other intensive sites. Indeed, the difficulty of scaling up may be greatest for ecological phenomena that are sensitive to historical disturbance and that exhibit the greatest spatiotemporal variation, such as denitrification in soils and the dynamics of bird communities. Our research shows that end member sites for some processes often provide important insights into the behavior of inherently heterogeneous ecological processes. In the current era of rapid environmental and biological change, key ecological responses at intensive sites will reflect both specific local drivers and regional trends.

Changes in autumn senescence in northern hemisphere deciduous trees: a meta-analysis of autumn phenology studies.

BACKGROUND AND AIMS: Many individual studies have shown that the timing of leaf senescence in boreal and temperate deciduous forests in the northern hemisphere is influenced by rising temperatures, but there is limited consensus on the magnitude, direction and spatial extent of this relationship. METHODS: A meta-analysis was conducted of published studies from the peer-reviewed literature that reported autumn senescence dates for deciduous trees in the northern hemisphere, encompassing 64 publications with observations ranging from 1931 to 2010. KEY RESULTS: Among the meteorological measurements examined, October temperatures were the strongest predictors of date of senescence, followed by cooling degree-days, latitude, photoperiod and, lastly, total monthly precipitation, although the strength of the relationships differed between high- and low-latitude sites. Autumn leaf senescence has been significantly more delayed at low (25° to 49°N) than high (50° to 70°N) latitudes across the northern hemisphere, with senescence across high-latitude sites more sensitive to the effects of photoperiod and low-latitude sites more sensitive to the effects of temperature. Delays in leaf senescence over time were stronger in North America compared with Europe and Asia. CONCLUSIONS: The results indicate that leaf senescence has been delayed over time and in response to temperature, although low-latitude sites show significantly stronger delays in senescence over time than high-latitude sites. While temperature alone may be a reasonable predictor of the date of leaf senescence when examining a broad suite of sites, it is important to consider that temperature-induced changes in senescence at high-latitude sites are likely to be constrained by the influence of photoperiod. Ecosystem-level differences in the mechanisms that control the timing of leaf senescence may affect both plant community interactions and ecosystem carbon storage as global temperatures increase over the next century.

Effects of precipitation regime and soil nitrogen on leaf traits in seasonally dry tropical forests of the Yucatan Peninsula, Mexico.

Leaf traits are closely associated with nutrient use by plants and can be utilized as a proxy for nutrient cycling processes. However, open questions remain, in particular regarding the variability of leaf traits within and across seasonally dry tropical forests. To address this, we considered six leaf traits (specific area, thickness, dry matter content, N content, P content and natural abundance (15)N) of four co-occurring tree species (two that are not associated with N2-fixing bacteria and two that are associated with N2-fixing bacteria) and net N mineralization rates and inorganic N concentrations along a precipitation gradient (537-1036 mm per year) in the Yucatan Peninsula, Mexico. Specifically we sought to test the hypothesis that leaf traits of dominant plant species shift along a precipitation gradient, but are affected by soil N cycling. Although variation among different species within each site explains some leaf trait variation, there is also a high level of variability across sites, suggesting that factors other than precipitation regime more strongly influence leaf traits. Principal component analyses indicated that across sites and tree species, covariation in leaf traits is an indicator of soil N availability. Patterns of natural abundance (15)N in foliage and foliage minus soil suggest that variation in precipitation regime drives a shift in plant N acquisition and the openness of the N cycle. Overall, our study shows that both plant species and site are important determinants of leaf traits, and that the leaf trait spectrum is correlated with soil N cycling.

Atmospheric inputs and nitrogen saturation status in and adjacent to Class I wilderness areas of the northeastern US.

Atmospheric inputs of N and S in bulk deposition (open collectors) and throughfall (beneath canopy collectors) were measured in and adjacent to two Class 1 wilderness areas of the northeastern US. In general, atmospheric S inputs followed our expectations with throughfall S fluxes increasing with elevation in the White Mountains, New Hampshire and throughfall S fluxes being greater in coniferous than deciduous stands in both sites. In contrast, throughfall N fluxes decreased significantly with elevation. Throughfall NO3 (-) fluxes were greater in coniferous than deciduous stands of Lye Brook, Vermont, but were greater in deciduous than coniferous stands of the White Mountains. We found overlap in the range of values for atmospheric N inputs between our measurements and monitoring data [National Atmospheric Deposition Program (NADP) and Clean Air Status and Trends Network (CASTNET)] for wet and total (wet + dry) deposition at Lye Brook. However, our measurements of total S deposition in the White Mountains and bulk (wet) deposition at both Lye Brook and the White Mountains were significantly lower than NADP plus CASTNET, and NADP data, respectively. Natural abundance (18)O in throughfall and bulk deposition were not significantly different, suggesting that there was no significant biological production of [Formula: see text] via nitrification in the canopy. NO3 (-) concentrations in streams were low and had natural abundance (18)O values consistent with microbial production, demonstrating that atmospheric N is being biologically transformed while moving through these watersheds and that these forested watersheds are unlikely to be N saturated.

Increased nitrogen leaching following soil freezing is due to decreased root uptake in a northern hardwood forest.

The depth and duration of snow pack is declining in the northeastern United States as a result of warming air temperatures. Since snow insulates soil, a decreased snow pack can increase the frequency of soil freezing, which has been shown to have important biogeochemical implications. One of the most notable effects of soil freezing is increased inorganic nitrogen losses from soil during the following growing season. Decreased nitrogen retention is thought to be due to reduced root uptake, but has not yet been measured directly. We conducted a 2-year snow-removal experiment at Hubbard Brook Experimental Forest in New Hampshire, USA to determine the effects of soil freezing on root uptake and leaching of inorganic nitrogen simultaneously. Snow removal significantly increased the depth of maximal soil frost by 37.2 and 39.5 cm in the first and second winters, respectively (P < 0.001 in 2008/2009 and 2009/2010). As a consequence of soil freezing, root uptake of ammonium declined significantly during the first and second growing seasons after snow removal (P = 0.023 for 2009 and P = 0.005 for 2010). These observed reductions in root nitrogen uptake coincided with significant increases in soil solution concentrations of ammonium in the Oa horizon (P = 0.001 for 2009 and 2010) and nitrate in the B horizon (P < 0.001 and P = 0.003 for 2009 and 2010, respectively). The excess flux of dissolved inorganic nitrogen from the Oa horizon that was attributable to soil freezing was 7.0 and 2.8 kg N ha(-1) in 2009 and 2010, respectively. The excess flux of dissolved inorganic nitrogen from the B horizon was lower, amounting to 1.7 and 0.7 kg N ha(-1) in 2009 and 2010, respectively. Results of this study provide direct evidence that soil freezing reduces root nitrogen uptake, demonstrating that the effects of winter climate change on root function has significant consequences for nitrogen retention and loss in forest ecosystems.

Winter climate change affects growing-season soil microbial biomass and activity in northern hardwood forests.

Understanding the responses of terrestrial ecosystems to global change remains a major challenge of ecological research. We exploited a natural elevation gradient in a northern hardwood forest to determine how reductions in snow accumulation, expected with climate change, directly affect dynamics of soil winter frost, and indirectly soil microbial biomass and activity during the growing season. Soils from lower elevation plots, which accumulated less snow and experienced more soil temperature variability during the winter (and likely more freeze/thaw events), had less extractable inorganic nitrogen (N), lower rates of microbial N production via potential net N mineralization and nitrification, and higher potential microbial respiration during the growing season. Potential nitrate production rates during the growing season were particularly sensitive to changes in winter snow pack accumulation and winter soil temperature variability, especially in spring. Effects of elevation and winter conditions on N transformation rates differed from those on potential microbial respiration, suggesting that N-related processes might respond differently to winter climate change in northern hardwood forests than C-related processes.

Global patterns of nitrogen limitation: confronting two global biogeochemical models with observations.

Projections of future changes in land carbon (C) storage using biogeochemical models depend on accurately modeling the interactions between the C and nitrogen (N) cycles. Here, we present a framework for analyzing N limitation in global biogeochemical models to explore how C-N interactions of current models compare to field observations, identify the processes causing model divergence, and identify future observation and experiment needs. We used a set of N-fertilization simulations from two global biogeochemical models (CLM-CN and O-CN) that use different approaches to modeling C-N interactions. On the global scale, net primary productivity (NPP) in the CLM-CN model was substantially more responsive to N fertilization than in the O-CN model. The most striking difference between the two models occurred for humid tropical forests, where the CLM-CN simulated a 62% increase in NPP at high N addition levels (30 g N m(-2) yr(-1)), while the O-CN predicted a 2% decrease in NPP due to N fertilization increasing plant respiration more than photosynthesis. Across 35 temperate and boreal forest sites with field N-fertilization experiments, we show that the CLM-CN simulated a 46% increase in aboveground NPP in response to N, which exceeded the observed increase of 25%. In contrast, the O-CN only simulated a 6% increase in aboveground NPP at the N-fertilization sites. Despite the small response of NPP to N fertilization, the O-CN model accurately simulated ecosystem retention of N and the fate of added N to vegetation when compared to empirical (15) N tracer application studies. In contrast, the CLM-CN predicted lower total ecosystem N retention and partitioned more losses to volatilization than estimated from observed N budgets of small catchments. These results point to the need for model improvements in both models in order to enhance the accuracy with which global C-N cycle feedbacks are simulated.

Influence of experimental snow removal on root and canopy physiology of sugar maple trees in a northern hardwood forest.

Due to projected increases in winter air temperatures in the northeastern USA over the next 100 years, the snowpack is expected to decrease in depth and duration, thereby increasing soil exposure to freezing air temperatures. To evaluate the potential physiological responses of sugar maple (Acer saccharum Marsh.) to a reduced snowpack, we measured root injury, foliar cation and carbohydrate concentrations, woody shoot carbohydrate levels, and terminal woody shoot lengths of trees in a snow manipulation experiment in New Hampshire, USA. Snow was removed from treatment plots for the first 6 weeks of winter for two consecutive years, resulting in lower soil temperatures to a depth of 50 cm for both winters compared to reference plots with an undisturbed snowpack. Visibly uninjured roots from trees in the snow removal plots had significantly higher (but sub-lethal) levels of relative electrolyte leakage than trees in the reference plots. Foliar calcium: aluminum (Al) molar ratios were significantly lower, and Al concentrations were significantly higher, in trees from snow removal plots than trees from reference plots. Snow removal also reduced terminal shoot growth and increased foliar starch concentrations. Our results are consistent with previous research implicating soil freezing as a cause of soil acidification that leads to soil cation imbalances, but are the first to show that this translates into altered foliar cation pools, and changes in soluble and structural carbon pools in trees. Increased soil freezing due to a reduced snowpack could exacerbate soil cation imbalances already caused by acidic deposition, and have widespread implications for forest health in the northeastern USA.

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.