Lars Vesterdal

Effects of thinning and soil properties on accumulation of carbon, nitrogen and phosphorus in the forest floor of Norway spruce stands

Abstract Area-based sampling was carried out to investigate the effect of thinning and soil properties on accumulation of forest floor carbon (C), nitrogen (N) and phosphorus (P) in Norway spruce (Picea abies (L.) Karst.) stands in Denmark. Four thinning intensities (unthinned, and about 83%, 67% and 50% of unthinned basal area) were investigated at three sites in Denmark: a calcareous, relatively nutrient rich soil with a sandy loam/loam texture and two soils with low to intermediate nutritional status and sandy loam and loamy sand textures, respectively. The effect of thinning on accumulated carbon and nitrogen was significant at two of the investigated sites. Accumulated phosphorus was significantly affected by thinning at one of these two sitesand at the third site. Accumulated carbon and phosphorus were negatively linearly correlated with thinning intensity. pH tended to be highest and C N and C P ratios tended to be lowest in the heaviest thinned plots. It is hypothesized that the differences in accumulation may be due to a more favourable microclimate and substrate for saprophytic organisms in the most heavily thinned plots. However, the differences between sites were greater than differences between thinning intensities. The accumulation of carbon, nitrogen and phosphorus in the forest floors was much higher at the two less fertile sites with loamy sand and sandy loam than at the relatively fertile site with sandy loam/loam. Significant differences in pH and in C N and C P ratios at the three sites indicate that the amounts of available nutrients influence the mineralization pattern. In addition, at the site with the greatest forest floor root density, competition for nutrients and moisture between mycorrhiza-infected roots and free-living saprophytic decomposers may be co-responsible for the large amounts of accumulated carbon, nitrogen and phosphorus.

Changes in soil properties after afforestation of former intensively managed soils with oak and Norway spruce

In many European countries, surplus agricultural production and ecological problems due to intensive soil cultivation have increased the interest in afforestation of arable soils. Many environmental consequences which might rise from this alternative land-use are only known from forest establishment on less intensively managed or marginal soils. The present study deals with changes in soil properties following afforestation of nutrient-rich arable soils. A chronosequence study was carried out comprising seven Norway spruce (Picea abies (Karst.) L.) and seven oak (Quercus robur L.) stands established from 1969 to 1997 on former horticultural and agricultural soils in the vicinity of Copenhagen, Denmark. For comparison, a permanent pasture and a ca. 200-year-old mixed deciduous forest were included. This paper reports on changes in pH values, base saturation (BSeff), exchangeable calcium, soil N pools (Nmin contents), and C/N ratios in the Ap-horizon (0–25 cm) and the accumulated forest floor. The results suggest that afforestation slowly modifies soil properties of former arable soils. Land-use history seems to influence soil properties more than the selected tree species. An effect of tree species was only found in the forest floor parameters. Soil acidification was the most apparent change along the chronosequence in terms of a pH decrease from 6 to 4 in the upper 5 cm soil. Forest floor pH varied only slightly around 5. Nitrogen storage in the Ap-horizon remained almost constant at 5.5 Mg N ha−1. This was less than in the mineral soil of the ca. 200-year-old forest. In the permanent pasture, N storage was somewhat higher in 0–15 cm depth than in afforested stands of comparable age. Nitrogen storage in the forest floor of the 0–30-year-old stands increased in connection with the build-up of forest floor mass. The increase was approximately five times greater under spruce than oak. Mineral soil C/N ratios ranged from 10 to 15 in all stands and tended to increase in older stands only in 0–5 cm depth. Forest floor C/N ratios were higher in spruce stands (26.4) as compared to oak stands (22.7). All stands except the youngest within a single tree species had comparable C/N ratios.

Influences of evergreen gymnosperm and deciduous angiosperm tree species on the functioning of temperate and boreal forests

It has been recognized for a long time that the overstorey composition of a forest partly determines its biological and physical–chemical functioning. Here, we review evidence of the influence of evergreen gymnosperm (EG) tree species and deciduous angiosperm (DA) tree species on the water balance, physical–chemical soil properties and biogeochemical cycling of carbon and nutrients. We used scientific publications based on experimental designs where all species grew on the same parent material and initial soil, and were similar in stage of stand development, former land use and current management. We present the current state of the art, define knowledge gaps, and briefly discuss how selection of tree species can be used to mitigate pollution or enhance accumulation of stable organic carbon in the soil. The presence of EGs generally induces a lower rate of precipitation input into the soil than DAs, resulting in drier soil conditions and lower water discharge. Soil temperature is generally not different, or slightly lower, under an EG canopy compared to a DA canopy. Chemical properties, such as soil pH, can also be significantly modified by taxonomic groups of tree species. Biomass production is usually similar or lower in DA stands than in stands of EGs. Aboveground production of dead organic matter appears to be of the same order of magnitude between tree species groups growing on the same site. Some DAs induce more rapid decomposition of litter than EGs because of the chemical properties of their tissues, higher soil moisture and favourable conditions for earthworms. Forest floors consequently tend to be thicker in EG forests compared to DA forests. Many factors, such as litter lignin content, influence litter decomposition and it is difficult to identify specific litter‐quality parameters that distinguish litter decomposition rates of EGs from DAs. Although it has been suggested that DAs can result in higher accumulation of soil carbon stocks, evidence from field studies does not show any obvious trend. Further research is required to clarify if accumulation of carbon in soils (i.e. forest floor + mineral soil) is different between the two types of trees. Production of belowground dead organic matter appears to be of similar magnitude in DA and EG forests, and root decomposition rate lower under EGs than DAs. However there are some discrepancies and still are insufficient data about belowground pools and processes that require further research. Relatively larger amounts of nutrients enter the soil–plant biogeochemical cycle under the influence of EGs than DAs, but recycling of nutrients appears to be slightly enhanced by DAs. Understanding the mechanisms underlying forest ecosystem functioning is essential to predicting the consequences of the expected tree species migration under global change. This knowledge can also be used as a mitigation tool regarding carbon sequestration or management of surface waters because the type of tree species affects forest growth, carbon, water and nutrient cycling.

Dramatic changes in ectomycorrhizal community composition, root tip abundance and mycelial production along a stand‐scale nitrogen deposition gradient

• Nitrogen (N) availability is known to influence ectomycorrhizal fungal components, such as fungal community composition, biomass of root tips and production of mycelia, but effects have never been demonstrated within the same forest. • We measured concurrently the abundance of ectomycorrhizal root tips and the production of external mycelia, and explored the changes in the ectomycorrhizal community composition, across a stand-scale N deposition gradient (from 27 to 43 kg N ha⁻¹ yr⁻¹) at the edge of a spruce forest. The N status was affected along the gradient as shown by a range of N availability indices. • Ectomycorrhizal root tip abundance and mycelial production decreased five and 10-fold, respectively, with increasing N deposition. In addition, the ectomycorrhizal fungal community changed and the species richness decreased. The changes were correlated with the measured indices of N status, in particular N deposition and N leaching. • The relationship between the altered ectomycorrhizal community, root tip abundance and mycelial production is discussed in the context of the N parameters. We suggest that increased N deposition to forests will cause large changes in ectomycorrhizal fungal community structure and functioning, which, in turn, may result in reduced N uptake by roots and fungi, and increased losses of N by leaching.

NITROGEN TURNOVER IN FOREST FLOORS OF COASTAL DOUGLAS‐FIR AT SITES DIFFERING IN SOIL NITROGEN CAPITAL

Nitrogen cycling is generally considered to be more rapid on sites with high availability of N; this is usually associated with differences in tree species composition. We tested whether N cycling in stands of a single tree species increased with increasing mineral soil nitrogen capital. Rates of N cycling in nine stands of Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) were estimated by measuring annual N input in litter, N content of the forest floor, and net N mineralization rate in the forest floor. Rates of C and N turnover were estimated from the litterfall:forest floor ratio. The amount of N returned in litter increased as soil N capital increased. The increase in litter N content resulted from both increased litter mass (C content) and increased N concentration in litter. Despite the greater litter input, forest floor mass was smaller at N-rich sites, indicating more rapid turnover of the forest floor on N-rich sites. There was a positive relationship between fractional annual loss of N from the forest floor and soil N capital. Therefore, even without changes in tree species composition, sites with greater soil N capital returned more N in annual litterfall and had faster turnover of N in the forest floor. Fractional annual loss of C also increased with increasing soil N capital, indicating faster decomposition on N-rich sites. The rate of net N mineralization during laboratory incubations of the forest floor was not correlated with soil N capital or N concentrations of litter, but was related to the C:N ratio of the forest floor. Net N mineralization was appreciable only at two sites where forest floor C:N ratios were <35. The rate of net N mineralization or C:N ratio of the forest floor were not good indicators of N availability at these sites. The results of this study are consistent with the hypothesis that rates of N cycling are faster on N-rich sites, and that this effect can occur in the absence of changes in tree species composition.

Soil carbon stock change following afforestation in Northern Europe: a meta‐analysis

Northern Europe supports large soil organic carbon (SOC) pools and has been subjected to high frequency of land-use changes during the past decades. However, this region has not been well represented in previous large-scale syntheses of land-use change effects on SOC, especially regarding effects of afforestation. Therefore, we conducted a meta-analysis of SOC stock change following afforestation in Northern Europe. Response ratios were calculated for forest floors and mineral soils (0-10 cm and 0-20/30 cm layers) based on paired control (former land use) and afforested plots. We analyzed the influence of forest age, former land-use, forest type, and soil textural class. Three major improvements were incorporated in the meta-analysis: analysis of major interaction groups, evaluation of the influence of nonindependence between samples according to study design, and mass correction. Former land use was a major factor contributing to changes in SOC after afforestation. In former croplands, SOC change differed between soil layers and was significantly positive (20%) in the 0-10 cm layer. Afforestation of former grasslands had a small negative (nonsignificant) effect indicating limited SOC change following this land-use change within the region. Forest floors enhanced the positive effects of afforestation on SOC, especially with conifers. Meta-estimates calculated for the periods <30 years and >30 years since afforestation revealed a shift from initial loss to later gain of SOC. The interaction group analysis indicated that meta-estimates in former land-use, forest type, and soil textural class alone were either offset or enhanced when confounding effects among variable classes were considered. Furthermore, effect sizes were slightly overestimated if sample dependence was not accounted for and if no mass correction was performed. We conclude that significant SOC sequestration in Northern Europe occurs after afforestation of croplands and not grasslands, and changes are small within a 30-year perspective.

Biotic homogenization can decrease landscape-scale forest multifunctionality

Significance Numerous studies have demonstrated the importance of biodiversity in maintaining multiple ecosystem functions and services (multifunctionality) at local spatial scales, but it is unknown whether similar relationships are found at larger spatial scales in real-world landscapes. Here, we show, for the first time to our knowledge, that biodiversity can also be important for multifunctionality at larger spatial scales in European forest landscapes. Both high local (α-) diversity and a high turnover in species composition between locations (high β-diversity) were found to be potentially important drivers of ecosystem multifunctionality. Our study provides evidence that it is important to conserve the landscape-scale biodiversity that is being eroded by biotic homogenization if ecosystem multifunctionality is to be maintained. Many experiments have shown that local biodiversity loss impairs the ability of ecosystems to maintain multiple ecosystem functions at high levels (multifunctionality). In contrast, the role of biodiversity in driving ecosystem multifunctionality at landscape scales remains unresolved. We used a comprehensive pan-European dataset, including 16 ecosystem functions measured in 209 forest plots across six European countries, and performed simulations to investigate how local plot-scale richness of tree species (α-diversity) and their turnover between plots (β-diversity) are related to landscape-scale multifunctionality. After accounting for variation in environmental conditions, we found that relationships between α-diversity and landscape-scale multifunctionality varied from positive to negative depending on the multifunctionality metric used. In contrast, when significant, relationships between β-diversity and landscape-scale multifunctionality were always positive, because a high spatial turnover in species composition was closely related to a high spatial turnover in functions that were supported at high levels. Our findings have major implications for forest management and indicate that biotic homogenization can have previously unrecognized and negative consequences for large-scale ecosystem multifunctionality.

Carbon Sequestration in Soil and Biomass Following Afforestation: Experiences from Oak and Norway Spruce Chronosequences in Denmark, Sweden and the Netherlands

There is limited knowledge of the contribution of afforested arable land to mitigation of greenhouse effects. In the AFFOREST project we evaluated the rate and magnitude of carbon (C) sequestration in biomass and soils following afforestation of cropland. Two oak (Quercus robur) and four Norway spruce (Picea abies) afforestation chronosequences (age range 1 to 90 years) were studied with respect to C sequestration in Denmark, Sweden and the Netherlands.

Carbon sequestration and forest management

Forest management has the potential to increase the terrestrial C pool. According to the rules of the Kyoto Protocol and of the United Nations Framework Convention on Climate Change, forestry can generate a sink for greenhouse gases that can contribute to meeting the national commitment to emissions reductions. Afforestation is a common strategy that over the course of decades leads to the incorporation of carbon dioxide (CO2) in plant biomass. However, site types such as wetlands and peatlands may even be a source of greenhouse gases when they are afforested. Adapted management of existing forests may have a less obvious or slower effect on the terrestrial C pool. It is mainly relevant in countries that already have a large forest cover. We analysed the effects of harvesting, rotation length, thinning, fertilizer application and tree-species selection. All these treatments have an impact on the forest productivity and consequently on C sequestration in the ecosystem. Many forest treatments are already an integral part of sustainable forestry practice. In the context of C sequestration and its accounting in national greenhouse-gas budgets, ecosystem stability is highly rated. Forests that are robust against disturbances up to a certain degree of severity are better suited for political commitments than stands of maximum productivity with a high risk of damages.

Distribution of biomass and carbon in even‐aged stands of Norway spruce (Picea abies (L.) Karst.): A case study on spacing and thinning effects in northern Denmark

The main objective of this case study was to explore the possible influence of forest management on the levels and distribution of biomass and carbon (C) in even-aged stands of Norway spruce [Picea abies (L.) Karst.] in Denmark. Data originated from a long-term thinning experiment and an adjacent spacing experiment at stand ages of 58 and 41 years, respectively. Biomass of 16 trees from different thinning and spacing treatments was measured or partly estimated, and soils were sampled for determination of C stocks. All trees in each plot were measured for stem diameter and some for total height, to allow for scaling-up results to stand-level estimates. For trees of similar size, foliage biomass tended to be higher in the spacing experiment, which was located on slightly more fertile land. Foliage biomass increased with increasing thinning grade, but the effect could not be separated from that of tree size. At stand level, foliage biomass tended to increase with increasing spacing as well as with increasing thinning grade. For branchwood, stems and roots (including below-ground stump), the biomass increased with increasing tree size and stand volume at tree and stand level, respectively, but no differences between stands, spacings or thinning grades were observed, apart from that expressed by tree size or stand volume. At stand level, C stocks of all biomass compartments decreased with increasing thinning grade, while the distribution between compartments was hardly influenced. The ratio between above-ground and stem biomass was about 1.21 at stand level, while the ratio between below- and above-ground biomass was about 0.17. Thinning influenced the C stock of the forest floor and mineral soil oppositely, resulting in no effect of thinning on total soil C.