Lucas E. Nave

Sustained carbon uptake and storage following moderate disturbance in a Great Lakes forest

Carbon (C) uptake rates in many forests are sustained, or decline only briefly, following disturbances that partially defoliate the canopy. The mechanisms supporting such functional resistance to moderate forest disturbance are largely unknown. We used a large-scale experiment, in which > 6700 Populus (aspen) and Betula (birch) trees were stem-girdled within a 39-ha area, to identify mechanisms sustaining C uptake through partial canopy defoliation. The Forest Accelerated Succession Experiment in northern Michigan, USA, employs a suite of C-cycling measurements within paired treatment and control meteorological flux tower footprints. We found that enhancement of canopy light-use efficiency and maintenance of light absorption maintained net ecosystem production (NEP) and aboveground wood net primary production (NPP) when leaf-area index (LAI) of the treatment forest temporarily declined by nearly half its maximum value. In the year following peak defoliation, redistribution of nitrogen (N) in the treatment forest from senescent early successional aspen and birch to non-girdled later successional species facilitated the recovery of total LAI to pre-disturbance levels. Sustained canopy physiological competency following disturbance coincided with a downward shift in maximum canopy height, indicating that compensatory photosynthetic C uptake by undisturbed, later successional subdominant and subcanopy vegetation supported C-uptake resistance to disturbance. These findings have implications for ecosystem management and modeling, demonstrating that forests may tolerate considerable leaf-area losses without diminishing rates of C uptake. We conclude that the resistance of C uptake to moderate disturbance depends not only on replacement of lost leaf area, but also on rapid compensatory photosynthetic C uptake during defoliation by emerging later successional species.

Fire effects on temperate forest soil C and N storage

Temperate forest soils store globally significant amounts of carbon (C) and nitrogen (N). Understanding how soil pools of these two elements change in response to disturbance and management is critical to maintaining ecosystem services such as forest productivity, greenhouse gas mitigation, and water resource protection. Fire is one of the principal disturbances acting on forest soil C and N storage and is also the subject of enormous management efforts. In the present article, we use meta-analysis to quantify fire effects on temperate forest soil C and N storage. Across a combined total of 468 soil C and N response ratios from 57 publications (concentrations and pool sizes), fire had significant overall effects on soil C (-26%) and soil N (-22%). The impacts of fire on forest floors were significantly different from its effects on mineral soils. Fires reduced forest floor C and N storage (pool sizes only) by an average of 59% and 50%, respectively, but the concentrations of these two elements did not change. Prescribed fires caused smaller reductions in forest floor C and N storage (-46% and -35%) than wildfires (-67% and -69%), and the presence of hardwoods also mitigated fire impacts. Burned forest floors recovered their C and N pools in an average of 128 and 103 years, respectively. Among mineral soils, there were no significant changes in C or N storage, but C and N concentrations declined significantly (-11% and -12%, respectively). Mineral soil C and N concentrations were significantly affected by fire type, with no change following prescribed burns, but significant reductions in response to wildfires. Geographic variation in fire effects on mineral soil C and N storage underscores the need for region-specific fire management plans, and the role of fire type in mediating C and N shifts (especially in the forest floor) indicates that averting wildfires through prescribed burning is desirable from a soils perspective.

Nitrogen Uptake by Trees and Mycorrhizal Fungi in a Successional Northern Temperate Forest: Insights from Multiple Isotopic Methods

Forest succession may cause changes in nitrogen (N) availability, vegetation and fungal community composition that affect N uptake by trees and their mycorrhizal symbionts. Understanding how these changes affect the functioning of the mycorrhizal symbiosis is of interest to ecosystem ecology because of the fundamental roles mycorrhizae play in providing nutrition to trees and structuring forest ecosystems. We investigated changes in tree and mycorrhizal fungal community composition, the availability and uptake of N by trees and mycorrhizal fungi in a forest undergoing a successional transition (age-related loss of early successional tree taxa). In this system, 82–96% of mycorrhizal hyphae were ectomycorrhizal (EM). As biomass production of arbuscular mycorrhizal (AM) trees increased, AM hyphae comprised a significantly greater proportion of total fungal hyphae, and the EM contribution to the N requirement of EM-associated tree taxa declined from greater than 75% to less than 60%. Increasing N availability was associated with lower EM hyphal foraging and 15N tracer uptake, yet the EM-associated later-successional species Quercus rubra was nonetheless a stronger competitor for 15N than AM-associated Acer rubrum, likely due to the more extensive nature of the persistent EM hyphal network. These results indicate that successional increases in N availability and co-dominance by AM-associated trees have increased the importance of AM fungi in the mycorrhizal community, while down-regulating EM N acquisition and transfer processes. This work advances understanding of linkages between tree and fungal community composition, and indicates that successional changes in N availability may affect competition between tree taxa with divergent resource acquisition strategies.

Changes in soil nitrogen cycling in a northern temperate forest ecosystem during succession

Nitrogen (N) transformations in forest soils are fundamentally important to plant and microbial N nutrition and the N balance of forest ecosystems, but changes in the patterns and rates of N transformations during forest succession are poorly understood. In order to better understand how soil N cycling changes during ecosystem succession, we analyzed four years of soil N cycling measurements in a 90-year-old secondary forest undergoing dieback of early-successional, dominant canopy trees. We expected that tree mortality would decrease root biomass, leading to increased soil NH4+ availability, and that these changes would prompt fundamental shifts in the N cycle such as the initiation of significant nitrification and increased cycling of oxidized N compounds in gas phase and soil solution. As expected, indices of soil NH4+ and NO3− availability increased with successional stage (defined as the proportion of dead trees), and were negatively correlated with the amount of fine root biomass. However, the standing amount of fine root biomass was not affected by tree mortality; increased soil NH4+ and NO3− availability therefore more likely resulted from successional increases in N-mineralization than decreases in root N uptake. Nitrification (as indicated by NO efflux as a proxy) increased due to elevated substrate (NH4+) availability, and the soil solution NO3− concentration increased as a result. Soil N2O efflux was not affected by succession, nor was it related to other N cycling parameters. Collectively, these results indicate that recent successional advancement has accelerated soil N cycling and shifted the N economy of this ecosystem towards greater importance of NO3−.

Networking our science to characterize the state, vulnerabilities, and management opportunities of soil organic matter

Soil organic matter (SOM) supports the Earths ability to sustain terrestrial ecosystems, provide food and fiber, and retains the largest pool of actively cycling carbon. Over 75% of the soil organic carbon (SOC) in the top meter of soil is directly affected by human land use. Large land areas have lost SOC as a result of land use practices, yet there are compensatory opportunities to enhance productivity and SOC storage in degraded lands through improved management practices. Large areas with and without intentional management are also being subjected to rapid changes in climate, making many SOC stocks vulnerable to losses by decomposition or disturbance. In order to quantify potential SOC losses or sequestration at field, regional, and global scales, measurements for detecting changes in SOC are needed. Such measurements and soil-management best practices should be based on well established and emerging scientific understanding of processes of C stabilization and destabilization over various timescales, soil types, and spatial scales. As newly engaged members of the International Soil Carbon Network, we have identified gaps in data, modeling, and communication that underscore the need for an open, shared network to frame and guide the study of SOM and SOC and their management for sustained production and climate regulation.

Reforestation can sequester two petagrams of carbon in US topsoils in a century

Significance Forestland in the United States is a carbon (C) sink, offsetting ∼10% of annual greenhouse gas emissions and mitigating climate change. Most of the C in forests is held in soils, and the capacity of forest soils to sequester C makes them a major component of the US forest C sink. Where reforestation is presently occurring, either through deliberate replanting after forestland is disturbed (e.g., burned), or where previously nonforested lands (e.g., cultivated) are converting to forestland, topsoils are accumulating C. However, these C accumulation rates are poorly constrained; quantifying them with empirical data are critical to accurately represent the role of reforestation in the US C budget and forecast the longevity of the US forest C sink. Soils are Earth’s largest terrestrial carbon (C) pool, and their responsiveness to land use and management make them appealing targets for strategies to enhance C sequestration. Numerous studies have identified practices that increase soil C, but their inferences are often based on limited data extrapolated over large areas. Here, we combine 15,000 observations from two national-level databases with remote sensing information to address the impacts of reforestation on the sequestration of C in topsoils (uppermost mineral soil horizons). We quantify C stocks in cultivated, reforesting, and natural forest topsoils; rates of C accumulation in reforesting topsoils; and their contribution to the US forest C sink. Our results indicate that reforestation increases topsoil C storage, and that reforesting lands, currently occupying >500,000 km2 in the United States, will sequester a cumulative 1.3–2.1 Pg C within a century (13–21 Tg C·y−1). Annually, these C gains constitute 10% of the US forest sector C sink and offset 1% of all US greenhouse gas emissions.

The role of reforestation in carbon sequestration

In the United States (U.S.), the maintenance of forest cover is a legal mandate for federally managed forest lands. More broadly, reforestation following harvesting, recent or historic disturbances can enhance numerous carbon (C)-based ecosystem services and functions. These include production of woody biomass for forest products, and mitigation of atmospheric CO2 pollution and climate change by sequestering C into ecosystem pools where it can be stored for long timescales. Nonetheless, a range of assessments and analyses indicate that reforestation in the U.S. lags behind its potential, with the continuation of ecosystem services and functions at risk if reforestation is not increased. In this context, there is need for multiple independent analyses that quantify the role of reforestation in C sequestration, from ecosystems up to regional and national levels. Here, we describe the methods and report the findings of a large-scale data synthesis aimed at four objectives: (1) estimate C storage in major ecosystem pools in forest and other land cover types; (2) quantify sources of variation in ecosystem C pools; (3) compare the impacts of reforestation and afforestation on C pools; (4) assess whether these results hold or diverge across ecoregions. The results of our synthesis support four overarching inferences regarding reforestation and other land use impacts on C sequestration. First, in the bigger picture, soils are the dominant C pool in all ecosystems and land cover types in the U.S., and soil C pool sizes vary less by land cover than by other factors, such as spatial variation or soil wetness. Second, where historically cultivated lands are being reforested, topsoils are sequestering significant amounts of C, with the majority of reforested lands yet to reach their capacity relative to the potential indicated by natural forest soils. Third, the establishment of woody vegetation delivers immediate to multi-decadal C sequestration benefits in aboveground woody biomass and coarse woody debris pools, with two- to three-fold C sequestration benefits in biomass during the first several decades following planting. Fourth, opportunities to enhance C sequestration through reforestation vary among the ecoregions, according to current levels of planting, typical forest growth rates, and past land uses (especially cultivation). Altogether, our results suggest that an immediate, but phased and spatially targeted approach to reforestation can enhance C sequestration in forest biomass and soils in the U.S. for decades to centuries to come.

Effects of canopy structure and species diversity on primary production in upper Great Lakes forests

Canopy structure and tree species diversity, shaped by succession, disturbance, and community composition, are linked to numerous ecosystem functions, including net primary production (NPP). Understanding of how ecosystem structural metrics are interrelated and mechanistically link to NPP, however, is incomplete. We characterized leaf area index (LAI), Simpson’s index of Diversity (D′, a measure of species diversity), and canopy rugosity (Rc, a measure of canopy physical complexity) in 11 forest stands comprising two chronosequences varying in establishing disturbance, and in three late successional communities. We related LAI, D′, and Rc to wood NPP (NPPw), and examined whether absorption of photosynthetically active radiation and light use-efficiency (LUE) link NPPw with ecosystem structure. We found that recovery of LAI and D′ was delayed following more severe establishing disturbances, but that the development of Rc was strikingly conserved regardless of disturbance, converging on a common mean value in late-successional stands irrespective of differences in leaf area index and species diversity. LAI was significantly correlated with NPPw in each stage of ecosystem development, but NPPw was only correlated with Rc in early successional stages and with D′ in late successional stages. Across all stands, NPPw was coupled with LAI and Rc, (but not D′) through positive relationships with light absorption and LUE. We conclude by advocating for better integration of ecological disciplines investigating structure–function interactions, suggesting that improved understanding of such relationships will require ecologists to traverse disciplinary boundaries.

Research Article: Soil respiration in upper Great Lakes old-growth forest ecosystems

Abstract. Forests, through photosynthetic fixation of carbon dioxide, serve as natural carbon (C) sinks, thereby offsetting a substantial fraction of anthropogenic greenhouse gas emissions. However, forests of the upper Great Lakes region are on average getting older, prompting uncertainty in the regions future capacity to sequester C. Several studies demonstrate higher-than-expected rates of C sequestration in biomass and soils, or net ecosystem production (NEP), of older forests, with declining C losses from soils a proposed mechanism sustaining NEP as forests age. Forest plots spanning nearly 200 years of ecosystem development and including three different old-growth forest types were used to examine changes with age in growing season soil respiration (Rs) - the C emitted from soils by roots and soil microbes. How heterotrophic soil respiration (Rh) - the C emitted from soils by microbes only - varies among old-growth forest types and with soil environment was also examined. Mean growing season Rs and soil temperature were significantly lower in old-growth forests, with the former declining as forests aged. Laboratory incubations and scaled estimates of Rh suggest significantly higher rates in more tree species diverse mixed canopy old-growth forests than in less diverse conifer and deciduous old-growth forest stands. Soil moisture was an important driver of Rh in all old-growth forest types, particularly in the top, carbon-rich organic horizon. Based on the results of this study, it can be concluded that declining C emissions from the soils of old-growth forests may contribute to unexpectedly high rates of forest C sequestration as forests age.