Calcium Additions and Microbial Nitrogen Cycle Processes in a Northern Hardwood Forest

Evaluating, and possibly ameliorating, the effects of base cation depletion in forest soils caused by acid deposition is an important topic in the northeastern United States. We added 850 kg Ca ha⁻¹ as wollastonite (CaSiO₃) to an 11.8-ha watershed at the Hubbard Brook Experimental Forest (HBEF), a northern hardwood forest in New Hampshire, USA, in fall 1999 to replace calcium (Ca) leached from the ecosystem by acid deposition over the past 6 decades. Soil microbial biomass carbon (C) and nitrogen (N) concentrations, gross and potential net N mineralization and nitrification rates, soil solution and stream chemistry, soil:atmosphere trace gas (CO₂, N₂O, CH₄) fluxes, and foliar N concentrations have been monitored in the treated watershed and in reference areas at the HBEF before and since the Ca addition. We expected that rates of microbial C and N cycle processes would increase in response to the treatment. By 2000, soil pH was increased by a full unit in the Oie soil horizon, and by 2002 it was increased by nearly 0.5 units in the Oa soil horizon. However, there were declines in the N content of the microbial biomass, potential net and gross N mineralization rates, and soil inorganic N pools in the Oie horizon of the treated watershed. Stream, soil solution, and foliar concentrations of N showed no response to treatment. The lack of stimulation of N cycling by Ca addition suggests that microbes may not be stimulated by increased pH and Ca levels in the naturally acidic soils at the HBEF, or that other factors (for example, phosphorus, or Ca binding of labile organic matter) may constrain the capacity of microbes to respond to increased pH in the treated watershed. Possible fates for the approximately 10 kg N ha⁻¹ decline in microbial and soil inorganic pools include components of the plant community that we did not measure (for example, seedlings, understory shrubs), increased fluxes of N₂ and/or N storage in soil organic matter. These results raise questions about the factors regulating microbial biomass and activity in northern hardwood forests that should be considered in the context of proposals to mitigate the depletion of nutrient cations in soil.     

Climate Variation and Soil Carbon and Nitrogen Cycling Processes in a Northern Hardwood Forest

We exploited the natural climate gradient in the northern hardwood forest at the Hubbard Brook Experimental Forest (HBEF) to evaluate the effects of climate variation similar to what is predicted to occur with global warming over the next 50–100 years for northeastern North America on soil carbon (C) and nitrogen (N) cycle processes. Our objectives were to (1) characterize differences in soil temperature, moisture and frost associated with elevation at the HBEF and (2) evaluate variation in total soil (TSR) and microbial respiration, N mineralization, nitrification, denitrification, nitrous oxide (N₂O) flux, and methane (CH₄) uptake along this gradient. Low elevation sites were consistently warmer (1.5–2.5°C) and drier than high elevation sites. Despite higher temperatures, low elevation plots had less snow and more soil frost than high elevation plots. Net N mineralization and nitrification were slower in warmer, low elevation plots, in both summer and winter. In summer, this pattern was driven by lower soil moisture in warmer soils and in winter the pattern was linked to less snow and more soil freezing in warmer soils. These data suggest that N cycling and supply to plants in northern hardwood ecosystems will be reduced in a warmer climate due to changes in both winter and summer conditions. TSR was consistently faster in the warmer, low elevation plots. N cycling processes appeared to be more sensitive to variation in soil moisture induced by climate variation, whereas C cycling processes appeared to be more strongly influenced by temperature.     

Impacts of Soil Nitrogen and Carbon Additions on Forest Understory Communities with a Long Nitrogen Deposition History

Rates of nitrogen (N) deposition have been historically high throughout much of the northeastern United States; thus, understanding the legacy of these high N loads is important for maintaining forest productivity and resilience. Though many studies have documented plant invasions due to N deposition and associated impacts on ecosystems, less is known about whether invasive plants will continue to increase in dominance with further shifting nutrient regimes. Using soil N and carbon additions, we examined the impact of both increasing and decreasing soil N on native and invasive understory plant dynamics over 4 years in a northeastern deciduous forest with a long history of N deposition. Despite applying large quantities of N, we found no difference in soil nitrate (NO₃) or ammonium (NH₄ ⁺) pools in N addition plots over the course of the study. Indicative of the potential N saturation in these forest soils, resin-available NO₃ ^? and NH₄ ⁺ showed evidence that the added N was rapidly moving out of the soil in N addition plots. Accordingly, we also found that adding N to soil altered neither invasive nor native plant abundance, though adding N temporally increased invasive plant richness. Carbon additions decreased soil N availability seasonally, but did not alter the total percent cover of invasive or native plants. Rather than being suppressed by excess N availability, native plant species in this ecosystem are primarily inhibited by the invasive species, which now dominate this site. In conclusion, understory plant communities in this potentially N-saturated ecosystem may be buffered to future alterations in N availability.     

Transport of Carbon and Nitrogen Between Litter and Soil Organic Matter in a Northern Hardwood Forest

We used sugar maple litter double-labeled with ¹³C and ¹⁵N to quantify fluxes of carbon (C) and nitrogen (N) between litter and soil in a northern hardwood forest and the retention of litter C and N in soil. Two cohorts of litter were compared, one in which the label was preferentially incorporated into non-structural tissue and the other structural tissue. Loss of ¹³C from this litter generally followed dry mass and total C loss whereas loss of ¹⁵N (20–30% in 1 year) was accompanied by large increases of total N content of this decaying litter (26–32%). Enrichment of ¹³C and ¹⁵N was detected in soil down to 10–15 cm depth. After 6 months of decay (November–May) 36–43% of the ¹³C released from the litter was recovered in the soil, with no differences between the structural and non-structural labeled litter. By October the percentage recovery of litter ¹³C in soil was much lower (16%). The C released from litter and remaining in soil organic matter (SOM) after 1 year represented over 30 g C m⁻² y⁻¹ of SOM accumulation. Recovery of litter ¹⁵N in soil was much higher than for C (over 90%) and in May ¹⁵N was mostly in organic horizons whereas by October it was mostly in 0–10 cm mineral soil. A small proportion of this N was recovered as inorganic N (2–6%). Recovery of ¹⁵N in microbial biomass was higher in May (13–15%) than in October (about 5%). The C:N ratio of the SOM and microbial biomass derived from the labeled litter was much higher for the structural than the non-structural litter and for the forest floor than mineral SOM, illustrating the interactive role of substrates and microbial activity in regulating the C:N stoichiometry of forest SOM formation. These results for a forest ecosystem long exposed to chronically high atmospheric N deposition (ca. 10 kg N ha⁻¹ y⁻¹) suggest possible mechanisms of N retention in soil: increased organic N leaching from fresh litter and reduced fungal transport of N from soil to decaying litter may promote N stabilization in mineral SOM even at a relatively low C:N ratio.     

Gross Nitrogen Dynamics in the Mycorrhizosphere of an Organic Forest Soil

The rhizosphere is a hot-spot for biogeochemical cycles, including production of greenhouse gases, as microbial activity is stimulated by rhizodeposits released by roots and mycorrhizae. The biogeochemical cycle of nitrogen (N) in soil is complex, consisting of many simultaneously occurring processes. In situ studies investigating the effects of roots and mycorrhizae on gross N turnover rates are scarce. We conducted a ¹⁵N tracer study under field conditions in a spruce forest on organic soil, which was subjected to exclusion of roots and roots plus ectomycorrhizae (ECM) for 6 years by trenching. The forest soil had, over the 6-year period, an average emission of nitrous oxide (N₂O) of 5.9 ± 2.1 kg N₂O ha^?1 year^?1. Exclusion of roots + ECM nearly tripled N₂O emissions over all years, whereas root exclusion stimulated N₂O emission only in the latest years and to a smaller extent. Gross mineralization–ammonium (NH₄ ⁺) immobilization turnover was enhanced by the presence of roots, probably due to high inputs of labile carbon, stimulating microbial activity. We found contrasting effects of roots and ECM on N₂O emission and mineralization, as the former was decreased but the latter was stimulated by roots and ECM. The N₂O emission was positively related to the ratio of gross NH₄ ⁺ oxidation (that is, autotrophic nitrification) to NH₄ ⁺ immobilization. Ammonium oxidation was only stimulated by the presence of ECM, but not by the presence of roots. Overall, we conclude that plants and their mycorrhizal symbionts actively control soil N cycling, thereby also affecting N₂O emissions from forest soils. Consequently, adapted forest management with permanent tree cover avoiding clearcutting could be a means to reduce N₂O emissions and potential N leaching; despite higher mineralization in the presence of roots and ECM, N₂O emissions are decreased as the relative importance of NH₄ ⁺ oxidation is decreased, mainly due to a stimulated microbial NH₄ ⁺ immobilization in the mycorrhizosphere.     

Thermal Properties of Tree Cavities During Winter in a Northern Hardwood Forest

ABSTRACT Tree cavities likely vary in their thermal quality for cavity-nesting animals, which could be especially important during winter. We conducted a winter field experiment to test whether cavities vary either in their buffering capacity or in their mean temperature according to predictable characteristics. We found that cavities buffered temperature and that there was a lag effect in temperature that appeared to be related to heating and cooling. Diameter at breast height was the most important variable influencing cavity temperature during the day, with smaller trees warming up more. During the night, diameter at breast height and tree decay class were important, such that larger, live trees cooled down less. Maintaining live trees with cavities in managed forests should be considered in addition to snag retention, because live trees appear to provide warmer structures during winter.     

Defining species guilds in the Central Hardwood Forest,USA

Tree regeneration outcomes are challenging to generalize and difficult to predict. Many tree species can establish new propagules in a variety of post-disturbance environments and many different reproductive mechanisms may be used. In order to develop conceptual models that accurately reflect reproductive potential, we need a better understanding of the similarities in regeneration ecology among species. We used information from the forest ecology literature to evaluate the reproductive attributes of sixty-two tree species in the central hardwood region of the eastern United States. Each species was classified categorically for features such as flowering, seed production and dispersal, seed dormancy, germination requirements, seedling characteristics, and vegetative reproduction. Cluster analysis (Jaccard's similarity coefficient, complete linkage method) and ordination (homogeneity analysis) were used to separate nine groups (guilds) of species that had similar reproductive attributes. Individual attributes that had high variance in the first and second dimensions included: seed banking, seed dispersal, seedling shade tolerance, and seedbed requirements. Members of each guild had similar levels of reproductive specialization and guilds were either pioneer-like, opportunistic, or persistent. Pioneer guilds included: short-lived or fugitive species that colonize sites rapidly and are too shade intolerant to replace themselves; shade-tolerant species that colonize frequently disturbed sites; and stress-tolerant pioneers that survive on dry or nutrient-poor sites. Opportunistic guilds contained species that are remarkably versatile in their reproductive effort. The most flexible opportunists can colonize new sites, maintain seed in a seed bank, sprout from existing stems and persist as a seedling or sapling bank. Persistent guilds contain species that develop and maintain advance regeneration. These include: species with moderate understory tolerance that regenerate via cycles of dieback and resprouting; and more tolerant species that maintain seedling or sapling banks. Our regeneration guilds may provide a useful approach for more realistically representing large and diverse sets of tree species in forest ecosystem models.     

Short-Term Effects of Fire and Other Fuel Reduction Treatments on Breeding Birds in a Southern Appalachian Upland Hardwood Forest

Abstract: We compared the effects of 3 fuel reduction techniques and a control on breeding birds during 2001-2005 using 50-m point counts. Four experimental units, each >14 ha, were contained within each of 3 replicate blocks at the Green River Game Land, Polk County, North Carolina, USA. Treatments were 1) prescribed burn, 2) mechanical understory reduction (chainsaw-felling of shrubs and small trees), 3) mechanical + burn, and 4) controls. We conducted mechanical treatments in winter 2001-2002 and prescribed burns in spring 2003. Tall shrub cover was substantially reduced in all treatments compared to controls. Tree mortality and canopy openness was highest in the mechanical + burn treatment after burning, likely due to higher fuel loading and hotter burns; tree mortality increased with time. Many bird species did not detectably decrease or increase in response to treatments. Species richness, total bird density, and some species, including indigo buntings (Passerina cyanea) and eastern bluebirds (Sialia sialis), increased in the mechanical + burn treatment after a 1-year to 2-year delay; eastern woodpewees (Contopus virens) increased immediately after treatment. Hooded warblers (Wilsonia citrina), black-and-white warblers (Mniotilta varia), and worm-eating warblers (Helmitheros vermivorus) declined temporarily in some or all treatments, likely in response to understory and (or) leaf litter depth reductions. Densities of most species affected by treatments varied with shrub cover, tree or snag density, or leaf litter depth. High snag availability, open conditions, and a higher density of flying insects in the mechanical + burn treatment likely contributed to increased bird density and species richness. In our study, fuel reduction treatments that left the canopy intact, such as low-intensity prescribed fire or mechanical understory removal, had few detectable effects on breeding birds compared to the mechanical + burn treatment. High-intensity burning with heavy tree-kill, as occurred in our mechanical + burn treatment, can be used as a management tool to increase densities of birds associated with open habitat while retaining many forest and generalist species, but may have short-term adverse effects on some species that are associated with the ground- or shrub-strata for nesting and foraging.     

Soil Solution Nitrogen Losses During Clearing of Lowland Amazon Forest for Pasture

Losses of nitrogen (N) often follow severe disturbance of forest ecosystems. In tropical forests, losses of N associated with the disturbance of clearing may be particularly important because rates of soil N cycling are high and forest clearing now occurs on a large scale. We measured soil solution inorganic N concentrations and fluxes for 1 year in an intact forest in the Brazilian Amazon state of Rondônia and in an adjacent 3-ha forest plot that was cleared for pasture by cutting, burning and planting pasture grass and in established cattle pastures on the same soils that were 5 and 22 years old. The cleared forest had higher soil solution NO ₃ ^? concentrations than the intact forest, but the difference between the cleared and control forests declined with time after the start of the first post-clearing rainy season. Established pastures had much lower solution NH ₄ ⁺ and NO ₃ ^? concentrations than forest or cleared forest. Estimated annual dissolved inorganic solution N fluxes to below 1 m during the first year after clearing were 2.5 kg ha^?1 in forest and 24.4 kg ha^?1 in newly cleared forest compared with only 0.5–1.2 kg ha^?1 in established pastures. The solution fluxes from cleared forest during the first year after clearing were approximately 7 times greater than gaseous N oxide (N₂O+NO) losses estimated for the same time. These results were consistent with the characterization of moist tropical forests on weathered soils as N-rich and likely to respond to disturbances that elevate soil N availability with increased loss to both soil solution and the atmosphere. These results also suggest that the relative increase in N oxide loss is substantially less than the increase solution inorganic N loss.     

Modeling the Effects of Harvest Alternatives on Mitigating Oak Decline in a Central Hardwood Forest Landscape

Oak decline is a process induced by complex interactions of predisposing factors, inciting factors, and contributing factors operating at tree, stand, and landscape scales. It has greatly altered species composition and stand structure in affected areas. Thinning, clearcutting, and group selection are widely adopted harvest alternatives for reducing forest vulnerability to oak decline by removing susceptible species and declining trees. However, the long-term, landscape-scale effects of these different harvest alternatives are not well studied because of the limited availability of experimental data. In this study, we applied a forest landscape model in combination with field studies to evaluate the effects of the three harvest alternatives on mitigating oak decline in a Central Hardwood Forest landscape. Results showed that the potential oak decline in high risk sites decreased strongly in the next five decades irrespective of harvest alternatives. This is because oak decline is a natural process and forest succession (e.g., high tree mortality resulting from intense competition) would eventually lead to the decrease in oak decline in this area. However, forest harvesting did play a role in mitigating oak decline and the effectiveness varied among the three harvest alternatives. The group selection and clearcutting alternatives were most effective in mitigating oak decline in the short and medium terms, respectively. The long-term effects of the three harvest alternatives on mitigating oak decline became less discernible as the role of succession increased. The thinning alternative had the highest biomass retention over time, followed by the group selection and clearcutting alternatives. The group selection alternative that balanced treatment effects and retaining biomass was the most viable alternative for managing oak decline. Insights from this study may be useful in developing effective and informed forest harvesting plans for managing oak decline.     

不同林窗马尾松凋落叶与土壤养分变化研究

以现有42年生的马尾松（Pinus massoniana）人工纯林,经过采伐形成4种不同大小有效面积的林窗（100、400、900和1 600 m²）为研究对象,以未经采伐的42年生马尾松人工纯林为对照样地,采用凋落叶分解袋法,研究不同大小有效面积林窗对马尾松凋落叶、土壤C、N、P及化学计量比和养分损失率的影响。研究结果表明：（1）不同大小有效面积林窗下的马尾松凋落叶、土壤C、N、P含量及养分损失率除土壤P含量和马尾松凋落叶P养分损失率外,均存在显著差异。随着林窗有效面积G1~G4的增大,马尾松凋落叶C、N、P含量均呈降低趋势,三者均在G3林窗体现出较小值。马尾松凋落叶C、N、P养分损失率、土壤C、N、P养分含量多呈抛物线趋势,且均在G2或G3林窗体现出最大值。（2）不同大小有效面积林窗下的马尾松凋落叶、土壤C/N/P均存在显著差异。随着林窗有效面积G1~G4的增大,马尾松人工林土壤C/N/P基本呈抛物线变化趋势,土壤C/N在G3林窗出现最大值,土壤C/P、N/P均在G2林窗体现出最大值;土壤C/N、C/P、N/P变异系数分别为13.31%、16.51%、17.21%。马尾松凋落叶C/N、C/P均在G3体现出最小值。（3）马尾松凋落叶C、N含量与土壤C、C/N/P及环境因子的相关性较强,P含量与它们的相关性较弱;C/N与土壤P、C/N/P及环境因子的相关性较强,C/P、N/P与土壤C/P及环境因子的相关性较强;C、N养分损失率与土壤C、C/N、C/P及环境因子的相关性较强,P养分损失率与土壤C、N、P含量及其化学计量比和环境因子的相关性较弱。土壤C、N、P含量及其化学计量比与环境因子的相关性较强。     

Spatial and Temporal Dynamics of Exotic Earthworm Communities Along Invasion Fronts in a Temperate Hardwood Forest in South-Central New York (USA)

The invasion of North American forests by exotic earthworms is producing profound ecosystem changes, such as alterations in soil nutrient cycling, and redistribution and loss of soil organic matter. However, the present and future extent of these invasions is difficult to evaluate without a better understanding of the factors that control the distribution and abundance of earthworms in previously non-invaded habitats. In this study, the species composition and short-term dynamics of three exotic earthworm invasion fronts were studied at a northern hardwood forest in south-central New York State (USA). Belt transects were established at each of the three locations to sample from earthworm-invaded areas through transition zones and into invasion front areas. Lumbricus rubellus, L. terrestrisandOctolasion tyrtaeum were the most common species, but their distribution was not homogeneous along the transects. Whereas, L. rubellus was the only species with relatively high adult densities at transition zones and invasion fronts, L. terrestris and O. tyrtaeum occurred mostly in the heavily earthworm-invaded areas and were rare at the invasion fronts. The density of earthworms along the transects decreased by 60–87 from June 2001 to October 2002 and then recovered in 2003 to values similar to those of 2001. This decrease was apparently caused by reduced recruitment of immature earthworms, probably related to the severe drought periods that the study area experienced in 2001 and 2002. Our data suggest that climate and topography, through their effects on soil moisture patterns, can be critical factors controlling the distribution and spread of exotic earthworms in previously non-invaded habitats.     

Impacts of Hemlock Loss on Nitrogen Retention Vary with Soil Nitrogen Availability in the Southern Appalachian Mountains

The impacts of exotic insects and pathogens on forest ecosystems are increasingly recognized, yet the factors influencing the magnitude of effects remain poorly understood. Eastern hemlock (Tsuga canadensis) exerts strong control on nitrogen (N) dynamics, and its loss due to infestation by the hemlock woolly adelgid (Adelges tsugae) is expected to decrease N retention in impacted stands. We evaluated the potential for site variation in N availability to influence the magnitude of effects of hemlock decline on N dynamics in mixed hardwood stands. We measured N pools and fluxes at three elevations (low, mid, high) subjected to increasing atmospheric N deposition where hemlock was declining or absent (as reference), in western North Carolina. Nitrogen pools and fluxes varied substantially with elevation and increasing N availability. Total forest floor and mineral soil N increased (P?<?0.0001, P?=?0.0017, resp.) and forest floor and soil carbon (C) to N ratio decreased with elevation (P?<?0.0001, P?=?0.0123, resp.), suggesting that these high elevation pools are accumulating available N. Contrary to expectations, subsurface leaching of inorganic N was minimal overall (<1?kg?ha^?1 9 months^?1), and was not higher in stands with hemlock mortality. Mean subsurface flux was 0.16?±?0.04 (SE) (kg?N?ha^?1 100?days^?1) in reference and 0.17?±?0.05 (kg?N?ha^?1 100?days^?1) in declining hemlock stands. Moreover, although subsurface N flux increased with N availability in reference stands, there was no relationship between N availability and flux in stands experiencing hemlock decline. Higher foliar N and observed increases in the growth of hardwood species in high elevation stands suggest that hemlock decline has stimulated N uptake and growth by healthy vegetation within this mixed forest, and may contribute to decoupling the relationship between N deposition and ecosystem N flux.     

Differences of Soil Microbial Biomass and Nitrogen Transformation under Two Forest Types in Central Germany

In order to observe the tree species effect on soil N status, soil microbial biomass C and N (C_mic, N_mic), potential N mineralization and potential nitrification (under laboratory incubation conditions, 22 °C) in different subhorizons (LOf₁, Of₂, Oh and mineral soil at 0–10 cm depth) were determined at three forest sites in central Germany. At each site, two contrasting stands (Beech, Norway spruce or Scots pine) were selected, where the initial soil conditions were similar. Three sampling dates that represented different stages of tree growth were selected: growing season - August, dormant season - November, after budbreak – April. In organic layers, C_mic-to-total C (C_t) ratios under beech and under conifer were 0.72–4.74% and 0.34–2.11%, respectively. N_mic-to-total N (N_t) ratios were 2.47–11.61% and 0.71–5.77%, respectively. Both concentrations of C_mic and N_mic were significantly affected by the stand type and sampling time. Potential N mineralization rates, ranging from 3.7 to 19.7 mg N kg⁻¹ d⁻¹, showed no clear pattern in relation to stand type. However, potential nitrification rates were mostly significantly higher under beech than under contrasting conifer. In mineral soils, concentrations of C_mic and N_mic showed a clear temporal pattern in the order: August>November>April. The average N_mic and N_mic-to-N_t were higher in soils from beech than conifer, while C_mic and C_mic-to-C_t ratios were similar between the two forest types. In organic layers, the highest values of C_mic-to-N_mic ratio and C_mic were found in November samples, especially under beech. By contrast, in mineral soils the highest value of C_mic-to-N_mic ratios were found in April samples, and at that time the C_mic concentrations were the lowest, especially under conifer. These results revealed the differences in microbial growth form and survival strategy associated with different tree species and soil layers.     

Fine Root Dynamics and Forest Production Across a Calcium Gradient in Northern Hardwood and Conifer Ecosystems

Losses of soil base cations due to acid rain have been implicated in declines of red spruce and sugar maple in the northeastern USA. We studied fine root and aboveground biomass and production in five northern hardwood and three conifer stands differing in soil Ca status at Sleepers River, VT; Hubbard Brook, NH; and Cone Pond, NH. Neither aboveground biomass and production nor belowground biomass were related to soil Ca or Ca:Al ratios across this gradient. Hardwood stands had 37% higher aboveground biomass (P = 0.03) and 44% higher leaf litter production (P < 0.01) than the conifer stands, on average. Fine root biomass (<2 mm in diameter) in the upper 35 cm of the soil, including the forest floor, was very similar in hardwoods and conifers (5.92 and 5.93 Mg ha⁻¹). The turnover coefficient (TC) of fine roots smaller than 1 mm ranged from 0.62 to 1.86 y⁻¹ and increased significantly with soil exchangeable Ca (P = 0.03). As a result, calculated fine root production was clearly higher in sites with higher soil Ca (P = 0.02). Fine root production (biomass times turnover) ranged from 1.2 to 3.7 Mg ha⁻¹ y⁻¹ for hardwood stands and from 0.9 to 2.3 Mg ha⁻¹ y⁻¹ for conifer stands. The relationship we observed between soil Ca availability and root production suggests that cation depletion might lead to reduced carbon allocation to roots in these ecosystems.     

Skid Trail Effects on Soil Methane and Carbon Dioxide Flux in a Selection-Managed Northern Hardwood Forest

Ecosystems - Temperate forest soils are an important sink for methane (CH4); however, disturbance through forest management and the creation of skid trails may significantly decrease soil’s...