Give and Take: A Watershed Acid Rain Mitigation Experiment Increases Baseflow Nitrogen Retention but Increases Stormflow Nitrogen Export.

In many temperate forested watersheds, hydrologic nitrogen export has declined substantially in recent decades, and many of these watersheds show enduring effects from historic acid deposition. A watershed acid remediation experiment in New Hampshire reversed many of these legacy effects of acid deposition and also increased watershed nitrogen export, suggesting that these two phenomena may be coupled. Here we examine stream nitrate dynamics in this watershed acid remediation experiment for indicators of nitrogen saturation in the terrestrial and aquatic ecosystems. Post-treatment, the (positive) slope of the relationship between nitrate concentration and discharge increased by a median of 82% ( p = 0.004). This resulted in greater flushing of nitrate during storm events, a key indicator of early stage nitrogen saturation. Hysteretic behavior of the concentration-discharge relationship indicated that the mobilization of soil nitrate pools was responsible for this increased flushing. In contrast to this evidence for nitrogen saturation in the terrestrial ecosystem, we found that nitrogen uptake increased, post-treatment, in the aquatic ecosystem, substantially attenuating growing-season nitrate flux by up to 71.1% ( p = 0.025). These results suggest that, as forests slowly recover from acid precipitation, terrestrial, and aquatic ecosystem nitrogen balance may be substantially altered.

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.

Forest biogeochemistry in response to drought.

Trees alter their use and allocation of nutrients in response to drought, and changes in soil nutrient cycling and trace gas flux (N2 O and CH4 ) are observed when experimental drought is imposed on forests. In extreme droughts, trees are increasingly susceptible to attack by pests and pathogens, which can lead to major changes in nutrient flux to the soil. Extreme droughts often lead to more common and more intense forest fires, causing dramatic changes in the nutrient storage and loss from forest ecosystems. Changes in the future manifestation of drought will affect carbon uptake and storage in forests, leading to feedbacks to the Earth's climate system. We must improve the recognition of drought in nature, our ability to manage our forests in the face of drought, and the parameterization of drought in earth system models for improved predictions of carbon uptake and storage in the world's 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.

Global environmental change and the nature of aboveground net primary productivity responses: insights from long-term experiments.

Many global change drivers chronically alter resource availability in terrestrial ecosystems. Such resource alterations are known to affect aboveground net primary production (ANPP) in the short term; however, it is unknown if patterns of response change through time. We examined the magnitude, direction, and pattern of ANPP responses to a wide range of global change drivers by compiling 73 datasets from long-term (>5 years) experiments that varied by ecosystem type, length of manipulation, and the type of manipulation. Chronic resource alterations resulted in a significant change in ANPP irrespective of ecosystem type, the length of the experiment, and the resource manipulated. However, the pattern of ecosystem response over time varied with ecosystem type and manipulation length. Continuous directional responses were the most common pattern observed in herbaceous-dominated ecosystems. Continuous directional responses also were frequently observed in longer-term experiments (>11 years) and were, in some cases, accompanied by large shifts in community composition. In contrast, stepped responses were common in forests and other ecosystems (salt marshes and dry valleys) and with nutrient manipulations. Our results suggest that the response of ANPP to chronic resource manipulations can be quite variable; however, responses persist once they occur, as few transient responses were observed. Shifts in plant community composition over time could be important determinants of patterns of terrestrial ecosystem sensitivity, but comparative, long-term studies are required to understand how and why ecosystems differ in their sensitivity to chronic resource alterations.

Root stress and nitrogen deposition: consequences and research priorities.

Stress within tree roots may influence whole-tree responses to nutrient deficiencies or toxic ion accumulation, but the mechanisms that govern root responses to the belowground chemical environment are poorly quantified. Currently, root production is modeled using rates of forest production and stoichiometry, but this approach alone may be insufficient to forecast variability in forest responses when physical and chemical stressors alter root lifespan, rooting depth or mycorrhizal colonization directly. Here, we review key research priorities for improving predictions of tree responses to changes in the belowground biogeochemical environment resulting from nitrogen deposition, including: limits of the optimum allocation paradigm, root physiological stress and lifespan, contingency effects that determine threshold responses across broad gradients, coupled water-biogeochemical interactions on roots, mycorrhizal dynamics that mediate root resilience and model frameworks to better simulate root feedbacks to aboveground function. We conclude that models incorporating physiological feedbacks, dynamic responses to coupled stressors, mycorrhizal interactions, and which challenge widely-accepted notions of optimum allocation, can elucidate potential thresholds of tree responses to biogeochemical stressors. Emphasis on comparative studies across species and environmental gradients, and which incorporates insights at the cellular and ecosystem level, is critical for forecasting whole-tree responses to altered biogeochemical landscapes.

Experimental approach and initial forest response to a simulated ice storm experiment in a northern hardwood forest.

Ice storms are a type of extreme winter weather event common to north temperate and boreal forests worldwide. Recent climate modelling studies suggest that these storms may become more frequent and severe under a changing climate. Compared to other types of storm events, relatively little is known about the direct and indirect impacts of these storms on forests, as naturally occurring ice storms are inherently difficult to study. Here we describe a novel experimental approach used to create a suite of ice storms in a mature hardwood forest in New Hampshire, USA. The experiment included five ice storm intensities (0, 6.4, 12.7 and 19.1 mm radial ice accretion) applied in a single year, and one ice storm intensity (12.7 mm) applied in two consecutive years. Results demonstrate the feasibility of this approach for creating experimental ice storms, quantify the increase in fine and coarse woody debris mass and nutrients transferred from the forest canopy to the soil under the different icing conditions, and show an increase in the damage to the forest canopy with increasing icing that evolves over time. In this forest, little damage occurred below 6.4 mm radial ice accretion, moderate damage occurred with up to 12.7 mm of accretion, and significant branch breakage and canopy damage occurred with 19.1 mm of ice. The icing in consecutive years demonstrated an interactive effect of ice storm frequency and severity such that some branches damaged in the first year of icing appeared to remain in the canopy and then fall to the ground in the second year of icing. These results have implications for National Weather Service ice storm warning levels, as they provide a quantitative assessment of ice-load related inputs of forest debris that will be useful to municipalities creating response plans for current and future ice storms.

Simulating Impacts of Ice Storms on Forest Ecosystems.

Ice storms can have profound and lasting effects on the structure and function of forest ecosystems in regions that experience freezing conditions. Current models suggest that the frequency and intensity of ice storms could increase over the coming decades in response to changes in climate, heightening interest in understanding their impacts. Because of the stochastic nature of ice storms and difficulties in predicting when and where they will occur, most past investigations of the ecological effects of ice storms have been based on case studies following major storms. Since intense ice storms are exceedingly rare events it is impractical to study them by waiting for their natural occurrence. Here we present a novel alternative experimental approach, involving the simulation of glaze ice events on forest plots under field conditions. With this method, water is pumped from a stream or lake and sprayed above the forest canopy when air temperatures are below freezing. The water rains down and freezes upon contact with cold surfaces. As the ice accumulates on trees, the boles and branches bend and break; damage that can be quantified through comparisons with untreated reference stands. The experimental approach described is advantageous because it enables control over the timing and amount of ice applied. Creating ice storms of different frequency and intensity makes it possible to identify critical ecological thresholds necessary for predicting and preparing for ice storm impacts.

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.

The response of terrestrial ecosystems to global climate change: towards an integrated approach.

Accumulating evidence points to an anthropogenic 'fingerprint' on the global climate change that has occurred in the last century. Climate change has, and will continue to have, profound effects on the structure and function of terrestrial ecosystems. As such, there is a critical need to continue to develop a sound scientific basis for national and international policies regulating carbon sequestration and greenhouse gas emissions. This paper reflects on the nature of current global change experiments, and provides recommendations for a unified multidisciplinary approach to future research in this dynamic field. These recommendations include: (1) better integration between experiments and models, and amongst experimental, monitoring, and space-for-time studies; (2) stable and increased support for long-term studies and multi-factor experiments; (3) explicit inclusion of biodiversity, disturbance, and extreme events in experiments and models; (4) consideration of timing vs intensity of global change factors in experiments and models; (5) evaluation of potential thresholds or ecosystem 'tipping points'; and (6) increased support for model-model and model-experiment comparisons. These recommendations, which reflect discussions within the TERACC international network of global change scientists, will facilitate the unraveling of the complex direct and indirect effects of global climate change on terrestrial ecosystems and their components.

Downsizing a long-term precipitation network: Using a quantitative approach to inform difficult decisions.

The design of a precipitation monitoring network must balance the demand for accurate estimates with the resources needed to build and maintain the network. If there are changes in the objectives of the monitoring or the availability of resources, network designs should be adjusted. At the Hubbard Brook Experimental Forest in New Hampshire, USA, precipitation has been monitored with a network established in 1955 that has grown to 23 gauges distributed across nine small catchments. This high sampling intensity allowed us to simulate reduced sampling schemes and thereby evaluate the effect of decommissioning gauges on the quality of precipitation estimates. We considered all possible scenarios of sampling intensity for the catchments on the south-facing slope (2047 combinations) and the north-facing slope (4095 combinations), from the current scenario with 11 or 12 gauges to only 1 gauge remaining. Gauge scenarios differed by as much as 6.0% from the best estimate (based on all the gauges), depending on the catchment, but 95% of the scenarios gave estimates within 2% of the long-term average annual precipitation. The insensitivity of precipitation estimates and the catchment fluxes that depend on them under many reduced monitoring scenarios allowed us to base our reduction decision on other factors such as technician safety, the time required for monitoring, and co-location with other hydrometeorological measurements (snow, air temperature). At Hubbard Brook, precipitation gauges could be reduced from 23 to 10 with a change of <2% in the long-term precipitation estimates. The decision-making approach illustrated in this case study is applicable to the redesign of monitoring networks when reduction of effort seems warranted.

Spruce-fir forest changes during a 30-year nitrogen saturation experiment.

A field experiment was established in a high elevation red spruce (Picea rubens Sarg.) - balsam fir (Abies balsamea) forest on Mount Ascutney Vermont, USA in 1988 to test the nitrogen (N) saturation hypothesis, and to better understand the mechanisms causing forest decline at the time. The study established replicate control, low and high dose nitrogen addition plots (i.e., 0, 15.7 and 31.4kgNH4Cl-Nha-1yr-1). The treatments began in 1988 and continued annually until 2010, but monitoring has continued to present. During the fertilization period, forest floor C:N, net in situ N mineralization, spruce foliar Ca%, and live spruce basal area decreased with increasing N addition, while foliar spruce N% and forest floor net nitrification increased with increasing N addition. The control plots aggraded forest floor N at a rate equal to the sum of the net in situ N mineralization plus average ambient deposition. Conversely, N addition plots lost forest floor N. Following the termination of N additions in 2010, the measured tree components returned to pre-treatment levels, but forest floor processes were slower to respond. During the 30year study, site surface air temperature has increased by 0.5°C per decade, and total N deposition has decreased 5.5 to 4.0kgNha-1yr-1. There have also been three significant drought years and at least one freeze injury year after which much of the forest mortality on the N addition plots occurred. Given that there was no control for the air temperature increase, discussion of the interactive impacts of climate and change and N addition is only subjective. Predicted changes in climate, N deposition and other stressors suggest that even in the absence of N saturation, regeneration of the spruce-fir ecosystem into the next century seems unlikely despite recent region-wide growth increases.

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.

Current status, uncertainty and future needs in soil organic carbon monitoring.

Increasing human demands on soil-derived ecosystem services requires reliable data on global soil resources for sustainable development. The soil organic carbon (SOC) pool is a key indicator of soil quality as it affects essential biological, chemical and physical soil functions such as nutrient cycling, pesticide and water retention, and soil structure maintenance. However, information on the SOC pool, and its temporal and spatial dynamics is unbalanced. Even in well-studied regions with a pronounced interest in environmental issues information on soil carbon (C) is inconsistent. Several activities for the compilation of global soil C data are under way. However, different approaches for soil sampling and chemical analyses make even regional comparisons highly uncertain. Often, the procedures used so far have not allowed the reliable estimation of the total SOC pool, partly because the available knowledge is focused on not clearly defined upper soil horizons and the contribution of subsoil to SOC stocks has been less considered. Even more difficult is quantifying SOC pool changes over time. SOC consists of variable amounts of labile and recalcitrant molecules of plant, and microbial and animal origin that are often operationally defined. A comprehensively active soil expert community needs to agree on protocols of soil surveying and lab procedures towards reliable SOC pool estimates. Already established long-term ecological research sites, where SOC changes are quantified and the underlying mechanisms are investigated, are potentially the backbones for regional, national, and international SOC monitoring programs.

Using ion-exchange resins to study soil response to experimental watershed acidification.

Ion-exchange resins (IER) offer alternative approaches to measuring ionic movement in soils that may have advantages over traditional approaches in some settings, but more information is needed to understand how IER compare with traditional methods of measurement in forested ecosystems. At the Bear Brook Watershed in Maine (BBWM), one of two paired, forested watersheds is treated bi-monthly with S and N (28.8 and 25.2kg ha(-1)yr(-1) of S and N, respectively). Both IER and ceramic cup tension lysimeters were used to study soil solution responses after approximately 11 years of treatment. Results from both methods showed treatments resulted in the mobilization of base cations and Al, and higher SO(4)-S and inorganic N in the treated watershed. Both methods indicated similar differences in results associated with forest type (hardwoods versus softwoods), a result of differences in litter quality and atmospheric aerosol interception capacity. The correlation between lysimeter and IER data for individual analytes varied greatly. Significant correlations were evident for Na (r=0.75), Al (r=0.65), Mn (r=0.61), Fe (r=0.57), Ca (r=0.49), K (r=0.41) and NO(3)-N (r=0.59). No correlation was evident between IER and soil solution data for NH(4)-N and Pb. Both IER and soil solution techniques suggested similar interpretations of biogeochemical behavior in the watershed.

Soilaluminum, iron, and phosphorus dynamics in response to long-term experimental nitrogen and sulfur additions at the Bear Brook Watershed in Maine, USA.

Atmospheric deposition of nitrogen (N) and sulfur (S) containing compounds affects soil chemistry in forested ecosystems through (1) acidification and the depletion of base cations, (2) metal mobilization, particularly aluminum (Al), and iron (Fe), (3) phosphorus (P) mobilization, and (4) N accumulation. The Bear Brook Watershed in Maine (BBWM) is a long-term paired whole-watershed experimental acidification study demonstrating evidence of each of these acidification characteristics in a northeastern U.S. forested ecosystem. In 2003, BBWM soils were studied using the Hedley fractionation procedure to better understand mechanisms of response in soil Al, Fe, and P chemistry. Soil P fractionation showed that recalcitrant P was the dominant fraction in these watersheds (49%), followed by Al and Fe associated P (24%), indicating that a majority of the soil P was biologically unavailable. Acidification induced mobilization of Al and Fe in these soils holds the potential for significant P mobilization. Forest type appears to exert important influences on metal and P dynamics. Soils supporting softwoods showed evidence of lower Al and Fe in the treated watershed, accompanied by lower soil P. Hardwood soils had higher P concentrations in surface soils as a result of increased biocycling in response to N additions in treatments. Accelerated P uptake and return in litterfall overshadowed acidification induced P mobilization and depletion mechanisms in hardwoods.