Climate and atmospheric deposition drive the inter-annual variability and long-term trend of dissolved organic carbon flux in the conterminous United States.

The lateral flux of dissolved organic carbon (DOC) from soils to inland waters and ultimately to the ocean represents a fundamental component of the global carbon cycle. To estimate the DOC flux, we developed an empirical terrestrial-aquatic DOC fluxes model (TAF-DOC). TAF-DOC incorporates various environmental factors (e.g., meteorology, sulfur, and nitrogen deposition) that to-date have not been comprehensively considered or well-represented in existing modeling frameworks. TAF-DOC was applied to estimate spatial-temporal patterns of DOC flux and potential fates across the conterminous United States during the 1985 to 2018 time period. Our results suggest that TAF-DOC successfully characterized spatial-temporal of DOC flux. As expected, the interannual pattern of DOC flux was strongly regulated by precipitation, but the long-term trend was significantly influenced by the rate of atmospheric wet sulfur deposition. From 1985 to 2018, TAF-DOC estimated DOC loading from terrestrial to aquatic ecosystems in the conterminous United States to be 33.5 ± 2.2 TgC per year, which was roughly 0.39-0.49% of total soil organic carbon stock estimates. The dominant fate of terrestrially-derived DOC was delivery to the coastal ocean in riverine export (41%), with another 21% buried in sediment and the remaining 12.8 ± 0.4 TgC per year (38%) returned to the atmosphere through outgassing from inland waters. Assuming the quantities of DOC sediment burial and export to the ocean as an annual sink of terrestrially-derived carbon, budget inventories and models that do not account for DOC flux in the conterminous United States will underestimate the net annual carbon sink by as much as 5.5-6.4%.

Forest density intensifies the importance of snowpack to growth in water-limited pine forests.

Warming climate and resulting declines in seasonal snowpack have been associated with drought stress and tree mortality in seasonally snow-covered watersheds worldwide. Meanwhile, increasing forest density has further exacerbated drought stress due to intensified tree-tree competition. Using a uniquely detailed data set of population-level forest growth (n = 2,495 sampled trees), we examined how inter-annual variability in growth relates to snow volume across a range of forest densities (e.g., competitive environments) in sites spanning a broad aridity gradient across the United States. Forest growth was positively related to snowpack in water-limited forests located at low latitude, and this relationship was intensified by forest density. However, forest growth was negatively related to snowpack in a higher latitude more energy-limited forest, and this relationship did not interact with forest density. Future reductions in snowpack may have contrasting consequences, as growth may respond positively in energy-limited forests and negatively in water-limited forests; however, these declines may be mitigated by reducing stand density through forest thinning.

Estimating uncertainty in the volume and carbon storage of downed coarse woody debris.

Downed coarse woody debris, also known as coarse woody detritus or downed dead wood, is challenging to estimate for many reasons, including irregular shapes, multiple stages of decay, and the difficulty of identifying species. In addition, some properties are commonly not measured, such as wood density and carbon concentration. As a result, there have been few previous evaluations of uncertainty in estimates of downed coarse woody debris, which are necessary for analysis and interpretation of the data. To address this shortcoming, we quantified uncertainties in estimates of downed coarse woody debris volume and carbon storage using data collected from permanent forest inventory plots in the northeastern United States by the Forest Inventory and Analysis program of the USDA Forest Service. Quality assurance data collected from blind remeasurement audits were used to quantify error in diameter measurements, hollowness of logs, species identification, and decay class determination. Uncertainty estimates for density, collapse ratio, and carbon concentration were taken from the literature. Estimates of individual sources of uncertainty were combined using Monte Carlo methods. Volume estimates were more reliable than carbon storage, with an average 95% confidence interval of 15.9 m3 /ha across the 79 plots evaluated, which was less than the mean of 31.2 m3 /ha. Estimates of carbon storage (and mass) were more uncertain, due to poorly constrained estimates of the density of wood. For carbon storage, the average 95% confidence interval was 11.1 Mg C/ha, which was larger than the mean of 4.6 Mg C/ha. Accounting for the collapse of dead wood as it decomposes would improve estimates of both volume and carbon storage. On the other hand, our analyses suggest that consideration of the hollowness of downed coarse woody debris pieces could be eliminated in this region, with little effect. This study demonstrates how uncertainty analysis can be used to quantify confidence in estimates and to help identify where best to allocate resources to improve monitoring designs.

The downed and dead wood inventory of forests in the United States.

The quantity and condition of downed dead wood (DDW) is emerging as a major factor governing forest ecosystem processes such as carbon cycling, fire behavior, and tree regeneration. Despite this, systematic inventories of DDW are sparse if not absent across major forest biomes. The Forest Inventory and Analysis program of the United States (US) Forest Service has conducted an annual DDW inventory on all coterminous US forest land since 2002 (~1 plot per 38,850 ha), with a sample intensification occurring since 2012 (~1 plot per 19,425 ha). The data are organized according to DDW components and by sampling information which can all be linked to a multitude of auxiliary information in the national database. As the sampling of DDW is conducted using field efficient line-intersect approaches, several assumptions are adopted during population estimation that serve to identify critical knowledge gaps. The plot- and population-level DDW datasets and estimates provide the first insights into an understudied but critical ecosystem component of temperate forests of North America with global application.

Patterns and drivers of recent disturbances across the temperate forest biome.

Increasing evidence indicates that forest disturbances are changing in response to global change, yet local variability in disturbance remains high. We quantified this considerable variability and analyzed whether recent disturbance episodes around the globe were consistently driven by climate, and if human influence modulates patterns of forest disturbance. We combined remote sensing data on recent (2001-2014) disturbances with in-depth local information for 50 protected landscapes and their surroundings across the temperate biome. Disturbance patterns are highly variable, and shaped by variation in disturbance agents and traits of prevailing tree species. However, high disturbance activity is consistently linked to warmer and drier than average conditions across the globe. Disturbances in protected areas are smaller and more complex in shape compared to their surroundings affected by human land use. This signal disappears in areas with high recent natural disturbance activity, underlining the potential of climate-mediated disturbance to transform forest landscapes.

Quantifying climate-growth relationships at the stand level in a mature mixed-species conifer forest.

A range of environmental factors regulate tree growth; however, climate is generally thought to most strongly influence year-to-year variability in growth. Numerous dendrochronological (tree-ring) studies have identified climate factors that influence year-to-year variability in growth for given tree species and location. However, traditional dendrochronology methods have limitations that prevent them from adequately assessing stand-level (as opposed to species-level) growth. We argue that stand-level growth analyses provide a more meaningful assessment of forest response to climate fluctuations, as well as the management options that may be employed to sustain forest productivity. Working in a mature, mixed-species stand at the Howland Research Forest of central Maine, USA, we used two alternatives to traditional dendrochronological analyses by (1) selecting trees for coring using a stratified (by size and species), random sampling method that ensures a representative sample of the stand, and (2) converting ring widths to biomass increments, which once summed, produced a representation of stand-level growth, while maintaining species identities or canopy position if needed. We then tested the relative influence of seasonal climate variables on year-to-year variability in the biomass increment using generalized least squares regression, while accounting for temporal autocorrelation. Our results indicate that stand-level growth responded most strongly to previous summer and current spring climate variables, resulting from a combination of individualistic climate responses occurring at the species- and canopy-position level. Our climate models were better fit to stand-level biomass increment than to species-level or canopy-position summaries. The relative growth responses (i.e., percent change) predicted from the most influential climate variables indicate stand-level growth varies less from to year-to-year than species-level or canopy-position growth responses. By assessing stand-level growth response to climate, we provide an alternative perspective on climate-growth relationships of forests, improving our understanding of forest growth dynamics under a fluctuating climate.

Drought timing and local climate determine the sensitivity of eastern temperate forests to drought.

Projected changes in temperature and drought regime are likely to reduce carbon (C) storage in forests, thereby amplifying rates of climate change. While such reductions are often presumed to be greatest in semi-arid forests that experience widespread tree mortality, the consequences of drought may also be important in temperate mesic forests of Eastern North America (ENA) if tree growth is significantly curtailed by drought. Investigations of the environmental conditions that determine drought sensitivity are critically needed to accurately predict ecosystem feedbacks to climate change. We matched site factors with the growth responses to drought of 10,753 trees across mesic forests of ENA, representing 24 species and 346 stands, to determine the broad-scale drivers of drought sensitivity for the dominant trees in ENA. Here we show that two factors-the timing of drought, and the atmospheric demand for water (i.e., local potential evapotranspiration; PET)-are stronger drivers of drought sensitivity than soil and stand characteristics. Drought-induced reductions in tree growth were greatest when the droughts occurred during early-season peaks in radial growth, especially for trees growing in the warmest, driest regions (i.e., highest PET). Further, mean species trait values (rooting depth and ψ50 ) were poor predictors of drought sensitivity, as intraspecific variation in sensitivity was equal to or greater than interspecific variation in 17 of 24 species. From a general circulation model ensemble, we find that future increases in early-season PET may exacerbate these effects, and potentially offset gains in C uptake and storage in ENA owing to other global change factors.

Large-scale disturbance legacies and the climate sensitivity of primary Picea abies forests.

Determining the drivers of shifting forest disturbance rates remains a pressing global change issue. Large-scale forest dynamics are commonly assumed to be climate driven, but appropriately scaled disturbance histories are rarely available to assess how disturbance legacies alter subsequent disturbance rates and the climate sensitivity of disturbance. We compiled multiple tree ring-based disturbance histories from primary Picea abies forest fragments distributed throughout five European landscapes spanning the Bohemian Forest and the Carpathian Mountains. The regional chronology includes 11,595 tree cores, with ring dates spanning the years 1750-2000, collected from 560 inventory plots in 37 stands distributed across a 1,000 km geographic gradient, amounting to the largest disturbance chronology yet constructed in Europe. Decadal disturbance rates varied significantly through time and declined after 1920, resulting in widespread increases in canopy tree age. Approximately 75% of current canopy area recruited prior to 1900. Long-term disturbance patterns were compared to an historical drought reconstruction, and further linked to spatial variation in stand structure and contemporary disturbance patterns derived from LANDSAT imagery. Historically, decadal Palmer drought severity index minima corresponded to higher rates of canopy removal. The severity of contemporary disturbances increased with each stand's estimated time since last major disturbance, increased with mean diameter, and declined with increasing within-stand structural variability. Reconstructed spatial patterns suggest that high small-scale structural variability has historically acted to reduce large-scale susceptibility and climate sensitivity of disturbance. Reduced disturbance rates since 1920, a potential legacy of high 19th century disturbance rates, have contributed to a recent region-wide increase in disturbance susceptibility. Increasingly common high-severity disturbances throughout primary Picea forests of Central Europe should be reinterpreted in light of both legacy effects (resulting in increased susceptibility) and climate change (resulting in increased exposure to extreme events).

Effects of thinning on drought vulnerability and climate response in north temperate forest ecosystems.

Reducing tree densities through silvicultural thinning has been widely advocated as a strategy for enhancing resistance and resilience to drought, yet few empirical evaluations of this approach exist. We examined detailed dendrochronological data from a long-term (> 50 years) replicated thinning experiment to determine if density reductions conferred greater resistance and/or resilience to droughts, assessed by the magnitude of stand-level growth reductions. Our results suggest that thinning generally enhanced drought resistance and resilience; however, this relationship showed a pronounced reversal over time in stands maintained at lower tree densities. Specifically, lower-density stands exhibited greater resistance and resilience at younger ages (49 years), yet exhibited lower resistance and resilience at older ages (76 years), relative to higher-density stands. We attribute this reversal to significantly greater tree sizes attained within the lower-density stands through stand development, which in turn increased tree-level water demand during the later droughts. Results from response-function analyses indicate that thinning altered growth-climate relationships, such that higher-density stands were more sensitive to growing-season precipitation relative to lower-density stands. These results confirm the potential of density management to moderate drought impacts on growth, and they highlight the importance of accounting for stand structure when predicting climate-change impacts to forests.

Singular and combined effects of blowdown, salvage logging, and wildfire on forest floor and soil mercury pools.

A number of factors influence the amount of mercury (Hg) in forest floors and soils, including deposition, volatile emission, leaching, and disturbances such as fire. Currently the impact on soil Hg pools from other widespread forest disturbances such as blowdown and management practices like salvage logging are unknown. Moreover, ecological and biogeochemical responses to disturbances are generally investigated within a single-disturbance context, with little currently known about the impact of multiple disturbances occurring in rapid succession. In this study we capitalize on a combination of blowdown, salvage logging and fire events in the sub-boreal region of northern Minnesota to assess both the singular and combined effects of these disturbances on forest floor and soil total Hg concentrations and pools. Although none of the disturbance combinations affected Hg in mineral soil, we did observe significant effects on both Hg concentrations and pools in the forest floor. Blowdown increased the mean Hg pool in the forest floor by 0.76 mg Hg m(-2) (223%). Salvage logging following blowdown created conditions leading to a significantly more severe forest floor burn during wildfire, which significantly enhanced Hg emission. This sequence of combined events resulted in a mean loss of approximately 0.42 mg Hg m(-2) (68% of pool) from the forest floor, after conservatively accounting for potential losses via enhanced soil leaching and volatile emissions between the disturbance and sampling dates. Fire alone or blowdown followed by fire did not significantly affect the total Hg concentrations or pools in the forest floor. Overall, unexpected consequences for soil Hg accumulation and by extension, atmospheric Hg emission and risk to aquatic biota, may result when combined impacts are considered in addition to singular forest floor and soil disturbances.

Carbon stocks across a chronosequence of thinned and unmanaged red pine (Pinus resinosa) stands.

Forests function as a major global C sink, and forest management strategies that maximize C stocks offer one possible means of mitigating the impacts of increasing anthropogenic CO2 emissions. We studied the effects of thinning, a common management technique in many forest types, on age-related trends in C stocks using a chronosequence of thinned and unmanaged red pine (Pinus resinosa) stands ranging from 9 to 306 years old. Live tree C stocks increased with age to a maximum near the middle of the chronosequence in unmanaged stands, and increased across the entire chronosequence in thinned stands. C in live understory vegetation and C in the mineral soil each declined rapidly with age in young stands but changed relatively little in middle-aged to older stands regardless of management. Forest floor C stocks increased with age in unmanaged stands, but forest floor C decreased with age after the onset of thinning around age 40 in thinned stands. Deadwood C was highly variable, but decreased with age in thinned stands. Total ecosystem C increased with stand age until approaching an asymptote around age 150. The increase in total ecosystem C was paralleled by an age-related increase in total aboveground C, but relatively little change in total belowground C. Thinning had surprisingly little impact on total ecosystem C stocks, but it did modestly alter age-related trends in total ecosystem C allocation between aboveground and belowground pools. In addition to characterizing the subtle differences in C dynamics between thinned and unmanaged stands, these results suggest that C accrual in red pine stands continues well beyond the 60-100 year management rotations typical for this system. Management plans that incorporate longer rotations and thinning in some stands could play an important role in maximizing C stocks in red pine forests while meeting other objectives including timber extraction, biodiversity conservation, restoration, and fuel reduction goals.

The efficacy of salvage logging in reducing subsequent fire severity in conifer-dominated forests of Minnesota, U.S.A.

Although primarily used to mitigate economic losses following disturbance, salvage logging has also been justified on the basis of reducing fire risk and fire severity; however, its ability to achieve these secondary objectives remains unclear. The patchiness resulting from a sequence of recent disturbances-blowdown, salvage logging, and wildfire-provided an excellent opportunity to assess the impacts of blowdown and salvage logging on wildfire severity. We used two fire-severity assessments (tree-crown and forest-floor characteristics) to compare post-wildfire conditions among three treatment combinations (Blowdown-Salvage-Fire, Blowdown-Fire, and Fire only). Our results suggest that salvage logging reduced the intensity (heat released) of the subsequent fire. However, its effect on severity (impact to the system) differed between the tree crowns and forest floor: tree-crown indices suggest that salvage logging decreased fire severity (albeit with modest statistical support), while forest-floor indices suggest that salvage logging increased fire severity. We attribute the latter finding to the greater exposure of mineral soil caused by logging operations; once exposed, soils are more likely to register the damaging effects of fire, even if fire intensity is not extreme. These results highlight the important distinction between fire intensity and severity when formulating post-disturbance management prescriptions.