Fungal biomass and ectomycorrhizal community assessment of phosphorus responsive Pinus taeda plantations.

Ectomycorrhizal fungi and non-ectomycorrhizal fungi are responsive to changes in environmental and nutrient availabilities. Although many species of ectomycorrhizas are known to enhance the uptake of phosphorus and other nutrients for Pinus taeda, it is not understood how to optimize these communities to have tangible effects on plantation silviculture and P use efficiency. The first step of this process is the identification of native fungi present in the system that are associated with P. taeda and influence P uptake efficiency. We used sand-filled mesh bags baited with finely ground apatite to sample ectomycorrhizal and non-ectomycorrhizal fungi associated with the rhizosphere of P-responsive P. taeda under several field conditions. Mesh bags were assessed for biomass accumulation over three years using a single three-month burial period pre-harvest and three six-month burial periods post-planting. Amplicon sequencing assessed ectomycorrhizal and non-ectomycorrhizal communities between phosphorus treatments, sites, mesh bags, and the rhizosphere of actively growing P. taeda in the field. We found biomass accumulation within the mesh bags was inversely related to increasing phosphorus fertilization (carryover) rates from pre-harvest to post-planting. Up to 25% increases in total biomass within the bags were observed for bags baited with P. Taxonomic richness was highest in Alfisol soils treated with phosphorus from the previous rotation and lowest in the Spodosol regardless of phosphorus treatment.

Moisture-driven divergence in mineral-associated soil carbon persistence.

Mineral stabilization of soil organic matter is an important regulator of the global carbon (C) cycle. However, the vulnerability of mineral-stabilized organic matter (OM) to climate change is currently unknown. We examined soil profiles from 34 sites across the conterminous USA to investigate how the abundance and persistence of mineral-associated organic C varied with climate at the continental scale. Using a novel combination of radiocarbon and molecular composition measurements, we show that the relationship between the abundance and persistence of mineral-associated organic matter (MAOM) appears to be driven by moisture availability. In wetter climates where precipitation exceeds evapotranspiration, excess moisture leads to deeper and more prolonged periods of wetness, creating conditions which favor greater root abundance and also allow for greater diffusion and interaction of inputs with MAOM. In these humid soils, mineral-associated soil organic C concentration and persistence are strongly linked, whereas this relationship is absent in drier climates. In arid soils, root abundance is lower, and interaction of inputs with mineral surfaces is limited by shallower and briefer periods of moisture, resulting in a disconnect between concentration and persistence. Data suggest a tipping point in the cycling of mineral-associated C at a climate threshold where precipitation equals evaporation. As climate patterns shift, our findings emphasize that divergence in the mechanisms of OM persistence associated with historical climate legacies need to be considered in process-based models.

Subterranean Survivorship and Seasonal Emergence of Laricobius spp. (Coleoptera: Derodontidae), Biological Control Agents for the Hemlock Woolly Adelgid.

Following the adventive arrival, subsequent spread, and ensuing impact of Adelges tsugae Annand (Hemiptera: Adelgidae), the hemlock woolly adelgid (HWA) in the eastern United States, a robust initiative was launched with the goal of decreasing ecosystem impacts from the loss of eastern hemlock (Pinales: Pinaceae). This initiative includes the use of biological control agents, including Laricobius spp. (Insecta: Coleoptera). Laboratory production of these agents is limited by subterranean mortality and early emergence. Therefore, the subterranean survivorship and timing of emergence of a mixture of Laricobius spp. was investigated. PVC traps internally lined with a sticky card and covered with a mesh screen were inserted into the soil to measure the percent emergence of adults based on the number of larvae placed within. The number of emerged adults in the field and laboratory-reared larval treatments was adjusted based on emergence numbers in the control and used as the response variable. Independent variables included in the final model were: treatment (field-collected vs. laboratory-reared), organic layer depth (cm), soil pH, and April-to-December mean soil moisture. No differences were found in survivorship between field-collected and laboratory-reared treatments. As pH and organic layer increased survivorship decreased, significantly. Although the majority of emergence occurred in the fall, emergence also occurred in spring and summer. The occurrence of spring and summer emergence and low survivorship (17.1 ± 0.4%) in the field across all treatments suggests that these are characteristics of Laricobius spp. field biology in their introduced range and not artifacts of the laboratory rearing process.

Climate Effects on Subsoil Carbon Loss Mediated by Soil Chemistry.

Subsoils store at least 50% of soil organic carbon (SOC) globally, but climate change may accelerate subsoil SOC (SOCsub) decomposition and amplify SOC-climate feedbacks. The climate sensitivity of SOCsub decomposition varies across systems, but we lack the mechanistic links needed to predict system-specific SOCsub vulnerability as a function of measurable properties at larger scales. Here, we show that soil chemical properties exert significant control over SOCsub decomposition under elevated temperature and moisture in subsoils collected across terrestrial National Ecological Observatory Network sites. Compared to a suite of soil and site-level variables, a divalent base cation-to-reactive metal gradient, linked to dominant mechanisms of SOCsub mineral protection, was the best predictor of the climate sensitivity of SOC decomposition. The response was "U"-shaped, showing higher sensitivity to temperature and moisture when either extractable base cations or reactive metals were highest. However, SOCsub in base cation-dominated subsoils was more sensitive to moisture than temperature, with the opposite relationship demonstrated in reactive metal-dominated subsoils. These observations highlight the importance of system-specific mechanisms of mineral stabilization in the prediction of SOCsub vulnerability to climate drivers. Our observations also form the basis for a spatially explicit, scalable, and mechanistically grounded tool for improved prediction of SOCsub response to climate change.

Emergent properties of microbial communities drive accelerated biogeochemical cycling in disturbed temperate forests.

Despite ever-increasing availability of detailed information about microbial community structure, relationships of microbial diversity with ecosystem functioning remain unclear. We investigated these relationships at the Coweeta Hydrologic Laboratory, where past forest disturbances (e.g., clear-cut) have altered both ecosystem processes (e.g., increased N export) and microbial communities (e.g., increased bacterial diversity). We sampled soils from disturbed and adjacent reference forests, characterized resident microbial communities, and measured several microbial C-cycle and N-cycle process rates. Microbial communities from historically disturbed soils exhibited altered ecosystem functioning, including generally higher rates of C- and N-cycle processes. Disturbed soil microbial communities also exhibited altered ecosystem multifunctionality, a composite variable consisting of all measured process rates as well as extracellular enzyme activities. Although we found few relationships between ecosystem functions and microbial alpha diversity, all functions were correlated with microbial community composition metrics, particularly r:K strategist ratios of bacterial phyla. Additionally, for both ecosystem multifunctionality and specific processes (i.e., C- and N-mineralization), microbial metrics significantly improved models seeking to explain variation in process rates. Our work sheds light on the links between microbial communities and ecosystem functioning and identifies specific microbial metrics important for modeling ecosystem responses to environmental change.

Soil Bacterial and Fungal Communities Exhibit Distinct Long-Term Responses to Disturbance in Temperate Forests.

In Appalachian ecosystems, forest disturbance has long-term effects on microbially driven biogeochemical processes such as nitrogen (N) cycling. However, little is known regarding long-term responses of forest soil microbial communities to disturbance in the region. We used 16S and ITS sequencing to characterize soil bacterial (16S) and fungal (ITS) communities across forested watersheds with a range of past disturbance regimes and adjacent reference forests at the Coweeta Hydrologic Laboratory in the Appalachian mountains of North Carolina. Bacterial communities in previously disturbed forests exhibited consistent responses, including increased alpha diversity and increased abundance of copiotrophic (e.g., Proteobacteria) and N-cycling (e.g., Nitrospirae) bacterial phyla. Fungal community composition also showed disturbance effects, particularly in mycorrhizal taxa. However, disturbance did not affect fungal alpha diversity, and disturbance effects were not consistent at the fungal class level. Co-occurrence networks constructed for bacteria and fungi showed that disturbed communities were characterized by more connected and tightly clustered network topologies, indicating that disturbance alters not only community composition but also potential ecological interactions among taxa. Although bacteria and fungi displayed different long-term responses to forest disturbance, our results demonstrate clear responses of important bacterial and fungal functional groups (e.g., nitrifying bacteria and mycorrhizal fungi), and suggest that both microbial groups play key roles in the long-term alterations to biogeochemical processes observed following forest disturbance in the region.

Soil Bacterial and Fungal Communities Show Distinct Recovery Patterns during Forest Ecosystem Restoration.

Bacteria and fungi are important mediators of biogeochemical processes and play essential roles in the establishment of plant communities, which makes knowledge about their recovery after extreme disturbances valuable for understanding ecosystem development. However, broad ecological differences between bacterial and fungal organisms, such as growth rates, stress tolerance, and substrate utilization, suggest they could follow distinct trajectories and show contrasting dynamics during recovery. In this study, we analyzed both the intra-annual variability and decade-scale recovery of bacterial and fungal communities in a chronosequence of reclaimed mined soils using next-generation sequencing to quantify their abundance, richness, ß-diversity, taxonomic composition, and cooccurrence network properties. Bacterial communities shifted gradually, with overlapping ß-diversity patterns across chronosequence ages, while shifts in fungal communities were more distinct among different ages. In addition, the magnitude of intra-annual variability in bacterial ß-diversity was comparable to the changes across decades of chronosequence age, while fungal communities changed minimally across months. Finally, the complexity of bacterial cooccurrence networks increased with chronosequence age, while fungal networks did not show clear age-related trends. We hypothesize that these contrasting dynamics of bacteria and fungi in the chronosequence result from (i) higher growth rates for bacteria, leading to higher intra-annual variability; (ii) higher tolerance to environmental changes for fungi; and (iii) stronger influence of vegetation on fungal communities.IMPORTANCE Both bacteria and fungi play essential roles in ecosystem functions, and information about their recovery after extreme disturbances is important for understanding whole-ecosystem development. Given their many differences in phenotype, phylogeny, and life history, a comparison of different bacterial and fungal recovery patterns improves the understanding of how different components of the soil microbiota respond to ecosystem recovery. In this study, we highlight key differences between soil bacteria and fungi during the restoration of reclaimed mine soils in the form of long-term diversity patterns, intra-annual variability, and potential interaction networks. Cooccurrence networks revealed increasingly complex bacterial community interactions during recovery, in contrast to much simpler and more isolated fungal network patterns. This study compares bacterial and fungal cooccurrence networks and reveals cooccurrences persisting through successional ages.

Microbial nitrogen cycling response to forest-based bioenergy production.

Concern over rising atmospheric CO2 and other greenhouse gases due to fossil fuel combustion has intensified research into carbon-neutral energy production. Approximately 15.8 million ha of pine plantations exist across the southeastern United States, representing a vast land area advantageous for bioenergy production without significant landuse change or diversion of agricultural resources from food production. Furthermore, intercropping of pine with bioenergy grasses could provide annually harvestable, lignocellulosic biomass feedstocks along with production of traditional wood products. Viability of such a system hinges in part on soil nitrogen (N) availability and effects of N competition between pines and grasses on ecosystem productivity. We investigated effects of intercropping loblolly pine (Pinus taeda) with switchgrass (Panicum virgatum) on microbial N cycling processes in the Lower Coastal Plain of North Carolina, USA. Soil samples were collected from bedded rows of pine and interbed space of two treatments, composed of either volunteer native woody and herbaceous vegetation (pine-native) or pure switchgrass (pine-switchgrass) in interbeds. An in vitro 15N pool-dilution technique was employed to quantify gross N transformations at two soil depths (0-5 and 5-15 cm) on four dates in 2012-2013. At the 0-5 cm depth in beds of the pine-switchgrass treatment, gross N mineralization was two to three times higher in November and February compared to the pine-native treatment, resulting in increased NH4(+) availability. Gross and net nitrification were also significantly higher in February in the same pine beds. In interbeds of the pine-switchgrass treatment, gross N mineralization was lower from April to November, but higher in February, potentially reflecting positive effects of switchgrass root-derived C inputs during dormancy on microbial activity. These findings indicate soil N cycling and availability has increased in pine beds of the pine-switchgrass treatment compared to those of the pine-native treatment, potentially alleviating any negative effects of N competition between pine and switchgrass. We expect that reduced soil C in the pine-switchgrass treatment, effects of pine and switchgrass rooting on soil C availability, and plant N demand are major factors influencing soil N transformations. Future research should examine rooting architecture in-intercropped systems and the effects on soil microbial communities and function.

Nitrous Oxide Fluxes in Fertilized L. Plantations across a Gradient of Soil Drainage Classes.

The effect of fertilizer management on nitrous oxide (NO) fluxes in agricultural ecosystems is well documented; however, our knowledge of these effects in managed forests is minimal. We established a comprehensive research study to address this knowledge gap across a range of soil drainage classes (poorly, moderately, and well drained) common in southern pine plantation management. Fertilizer treatments in each drainage class comprised of control (no fertilizer), urea + phosphorus (P), and P-coated urea fertilizer (CUF). Fertilization (168 kg N ha) occurred independently during the spring, summer, and fall to assess the effects of application timing. Nitrous oxide sampling, using vented static chambers, started immediately after seasonal fertilizer application and was performed every 6 wk for more than 1 yr. Time-integrated net annual NO emissions increased with urea (1.15 kg NO-N ha) and CUF (0.88 kg NO-N ha) application compared with unfertilized control (0.22 kg NO-N ha). Mean annual NO flux was significantly increased with fall fertilization (1.17 kg NO-N ha) relative to spring (0.73 kg NO-N ha) or summer (0.33 kg NO-N ha). Similarly, average annual NO flux was higher in poorly drained soils (1.40 kg NO-N ha) than in moderately drained (0.46 kg NO-N ha) and well-drained soils (0.39 kg NO-N ha). This study suggests that NO emissions after fertilization can be minimized by avoiding fall fertilization and poorly drained soils and by selecting enhanced-efficiency N fertilizers over urea.

Nitrogen leaching from Douglas-fir forests after urea fertilization.

Leaching of nitrogen (N) after forest fertilization has the potential to pollute ground and surface water. The purpose of this study was to quantify N leaching through the primary rooting zone of N-limited Douglas-fir [Pseudotsuga menziesii (Mirb.) Franco] forests the year after fertilization (224 kg N ha(-1) as urea) and to calculate changes in the N pools of the overstory trees, understory vegetation, and soil. At six sites on production forests in the Hood Canal watershed, Washington, tension lysimeters and estimates of the soil water flux were used to quantify the mobilization and leaching of NO(3)-N, NH(4)-N, and dissolved organic nitrogen below the observed rooting depth. Soil and vegetation samples were collected before fertilization and 1 and 6 mo after fertilization. In the year after fertilization, the total leaching beyond the primary rooting zone in excess of control plots was 4.2 kg N ha(-1) (p = 0.03), which was equal to 2% of the total N applied. The peak NO(3)-N concentration that leached beyond the rooting zone of fertilized plots was 0.2 mg NO(3)-N L(-1). Six months after fertilization, 26% of the applied N was accounted for in the overstory, and 27% was accounted for in the O+A horizon of the soil. The results of this study indicate that forest fertilization can lead to small N leaching fluxes out of the primary rooting zone during the first year after urea application.