Stephan Zimmermann

Critical examination of trace element enrichments and depletions in soils: As, Cr, Cu, Ni, Pb, and Zn in Swiss forest soils

The aim of this study was to obtain an overview of trace element concentrations in Swiss forest soils and to critically assess the measured values with respect to anthropogenic input vs. lithogenic background. Twenty-three sites were selected which represent a broad range of natural forest sites, bedrock material and soil types of Switzerland. At each site, samples were collected from all genetic soil horizons down to a C or B/C horizon. Total concentrations of As, Cr, Cu, Ni, Pb, and Zn in all samples were determined by X-ray fluorescence spectrometry. There were distinct differences in the geological background values estimated from the concentrations measured in the samples from the lowest soil horizon. Background concentrations for Cr and Ni were lowest in granite and gneiss, whereas Pb and Zn were highest in limestone and marl. Enrichment or depletion of the trace elements was assessed using Zr as reference element. Within the same profile, the six trace elements showed completely different enrichment/depletion patterns with depth. The various natural processes and anthropogenic inputs that can lead to these patterns are critically discussed. Based on this critical assessment, pollution of the investigated forest soils was found to be most severe for Pb and Zn and to a somewhat lesser extent for As and Cu, whereas anthropogenic input of Cr and Ni seems to be less important. The data suggest that a critical evaluation of enrichment factors is a better tool to assess soil pollution with trace elements than the use of maximum allowable concentrations (MAC) for topsoil samples. The enrichment factors calculated as described here consider the effects of geological variation on metal abundances whereas the MAC does not. In order to obtain an estimate of soil solution concentrations, water extracts of the samples collected from a subset of 10 soil profiles were analyzed for the same trace elements. Solubility of all elements generally decreased with soil depth. An exception was Cr, Cu, and Ni solubility in the humus layer, which was lower than in the underlying mineral horizon. For all elements, solubility was higher for the collective of soil samples depleted in this element when compared to the samples, in which the element was enriched.

Resistance and resilience of the forest soil microbiome to logging-associated compaction

Soil compaction is a major disturbance associated with logging, but we lack a fundamental understanding of how this affects the soil microbiome. We assessed the structural resistance and resilience of the microbiome using a high-throughput pyrosequencing approach in differently compacted soils at two forest sites and correlated these findings with changes in soil physical properties and functions. Alterations in soil porosity after compaction strongly limited the air and water conductivity. Compaction significantly reduced abundance, increased diversity, and persistently altered the structure of the microbiota. Fungi were less resistant and resilient than bacteria; clayey soils were less resistant and resilient than sandy soils. The strongest effects were observed in soils with unfavorable moisture conditions, where air and water conductivities dropped well below 10% of their initial value. Maximum impact was observed around 6–12 months after compaction, and microbial communities showed resilience in lightly but not in severely compacted soils 4 years post disturbance. Bacteria capable of anaerobic respiration, including sulfate, sulfur, and metal reducers of the Proteobacteria and Firmicutes, were significantly associated with compacted soils. Compaction detrimentally affected ectomycorrhizal species, whereas saprobic and parasitic fungi proportionally increased in compacted soils. Structural shifts in the microbiota were accompanied by significant changes in soil processes, resulting in reduced carbon dioxide, and increased methane and nitrous oxide emissions from compacted soils. This study demonstrates that physical soil disturbance during logging induces profound and long-lasting changes in the soil microbiome and associated soil functions, raising awareness regarding sustainable management of economically driven logging operations.

Morphological and physiological responses of Scots pine fine roots to water supply in a dry climatic region in Switzerland.

In recent decades, Scots pine (Pinus sylvestris L.) forests in inner-Alpine dry valleys of Switzerland have suffered from drought and elevated temperatures, resulting in a higher mortality rate of trees than the mean mortality rate in Switzerland. We investigated the responses of fine roots (standing crop, morphological and physiological features) to water supply in a Scots pine forest in the Rhone valley. Before irrigation started in 2003, low- and high-productivity Scots pine trees were selected based on their crown transparency. The fine root standing crop measured in spring from 2003 to 2005 was unaffected by the irrigation treatment. However, irrigation significantly enhanced the fine root standing crop during the vegetation period when values from spring were compared with values from fall in 2005. Irrigation slightly increased specific root length but decreased root tissue density. Fine root O2-consumption capacity decreased slightly in response to the irrigation treatment. Using ingrowth cores to observe the responses of newly produced fine roots, irrigation had a significantly positive effect on the length of fine roots, but there were no differences between the low- and high-productivity trees. In contrast to the weak response of fine roots to irrigation, the aboveground parts responded positively to irrigation with more dense crowns. The lack of a marked response of the fine root biomass to irrigation in the low- and high-productivity trees suggests that fine roots have a high priority for within-tree carbon allocation.

Heavy-Machinery Traffic Impacts Methane Emissions as Well as Methanogen Abundance and Community Structure in Oxic Forest Soils

ABSTRACT Temperate forest soils are usually efficient sinks for the greenhouse gas methane, at least in the absence of significant amounts of methanogens. We demonstrate here that trafficking with heavy harvesting machines caused a large reduction in CH4 consumption and even turned well-aerated forest soils into net methane sources. In addition to studying methane fluxes, we investigated the responses of methanogens after trafficking in two different forest sites. Trafficking generated wheel tracks with different impact (low, moderate, severe, and unaffected). We found that machine passes decreased the soils macropore space and lowered hydraulic conductivities in wheel tracks. Severely compacted soils yielded high methanogenic abundance, as demonstrated by quantitative PCR analyses of methyl coenzyme M reductase (mcrA) genes, whereas these sequences were undetectable in unaffected soils. Even after a year after traffic compression, methanogen abundance in compacted soils did not decline, indicating a stability of methanogens here over time. Compacted wheel tracks exhibited a relatively constant community structure, since we found several persisting mcrA sequence types continuously present at all sampling times. Phylogenetic analysis revealed a rather large methanogen diversity in the compacted soil, and most mcrA gene sequences were mostly similar to known sequences from wetlands. The majority of mcrA gene sequences belonged either to the order Methanosarcinales or Methanomicrobiales, whereas both sites were dominated by members of the families Methanomicrobiaceae Fencluster, with similar sequences obtained from peatland environments. The results show that compacting wet forest soils by heavy machinery causes increases in methane production and release.

Effects of Land-Use Change on Carbon Stocks in Switzerland

We assessed how consequences of future land-use change may affect size and spatial shifts of C stocks under three potential trends in policy—(a) business-as-usual: continuation of land-use trends observed during the past 15xa0years; (b) extensification: full extensification of open-land; and (c) liberalization: full reforestation potential. The build-up times for the three scenarios are estimated at 30, 80 and 100xa0years, respectively. Potential C-stock change rates are derived from the literature. Whereas the business-as-usual scenario would cause marginal changes of 0.5%, liberalization would provoke a 13% increase in C stocks (+62xa0MtC). Gains of 24% would be expected for forests (+95xa0MtC), whereas open-land C stock would decrease 27% (−33xa0MtC). Extensification would lead to a C stock decrease of 3% (−12xa0MtC). Whereas forest C is expected to increase 12% (+36.5xa0MtC) at high elevations, stocks of open-land C would decline 38.5% (−48.5xa0MtC). Most affected are unfavorable grasslands, which increase in area (+59%) but contribute only 14.5% to the C stocks. C sinks would amount to 0.6xa0MtCxa0y−1 assuming a build-up time of 100xa0years for the liberalization scenario. C stocks on the current forest area are increasing by 1xa0MtCxa0y−1. The maximal total C sink of 1.6xa0MtC might thus suffice to compensate for agricultural greenhouse gases (2004: 1.4xa0Mt CO2–C equivalents), but corresponds only to 11–13% of the anthropogenic greenhouse gas emission in Switzerland. Thus, even the largest of the expected terrestrial C stocks under liberalization will be small in comparison with current emissions of anthropogenic greenhouse gases.

Wood-ash recycling affects forest soil and tree fine-root chemistry and reverses soil acidification

Wood ash was applied to a forest ecosystem with the aim to recycle nutrients taken from the forest and to mitigate the negative effects of intensive harvesting. After two years, the application of 8,000 kg ha−1 of wood ash increased soil exchangeable Ca and Mg. Similarly, an increase in Ca and Mg in the Norway spruce fine roots was recorded, leading to significant linear correlations between soil and root Ca and soil and root Mg. In contrast to these macronutrients, the micronutrients Fe and Zn and the toxic element Al decreased in the soil exchangeable fraction with the addition of wood ash, but not in the fine roots. Only Mn decreased in soil and in fine roots leading to a significant linear correlation between soil and root Mn. In soil, as well as in fine roots, strong positive correlations were found between the elements Ca and Mg and between Fe and Al. This indicates that the uptake of Mg resembles that of Ca and that of Al that of Fe. With the wood ash application, the pH increased from 3.2 to 4.8, the base saturation from 30% to 86%, the molar basic cations/Al ratio (BC/Al) of the soil solution from 1.5 to 5.5, and the molar Ca/Al ratio of the fine roots from 1.3 to 3.7. Overall, all below-ground indicators of soil acidification responded positively to the wood ash application within two years. Nitrate concentrations increased only slightly in the soil solution at a soil depth of 75–80 cm, and no signs of increased heavy metal concentrations in the soils or in the fine roots were apparent. This suggests that the recycling of wood ash could be an integral part of sustainable forest management because it closes the nutrient cycle and reverses soil acidification.

Afforestation with Norway spruce on a subalpine pasture alters carbon dynamics but only moderately affects soil carbon storage

There is a strong trend toward reforestation of abandoned grasslands in alpine regions which may impact the carbon balance of alpine ecosystems. Here, we studied the effects of afforestation with Norway spruce (Picea abies L.) on an extensively grazed subalpine pasture in Switzerland on soil organic carbon (SOC) cycling and storage. Along a 120-year long chronosequence with spruce stands of 25, 30, 40, 45, and >120xa0years and adjacent pastures, we measured tree biomass, SOC stocks down to the bedrock, natural 13C abundances, and litter quality. To unravel controls on SOC cycling, we have monitored microclimatic conditions and quantified SOC decomposability under standardized conditions as well as soil respiration in situ. Stocks of SOC were only moderately affected by the afforestation: in the mineral soil, SOC stocks transiently decreased after tree establishment, reaching a minimum 40–45xa0years after afforestation (−25xa0%) and increased thereafter. Soils of the mature spruce forest stored the largest amount of SOC, 13xa0% more than the pasture soils, mainly due to the accumulation of an organic layer (23xa0txa0Cxa0ha−1). By comparison, C accumulated in the tree biomass exceeded the SOC pool by a factor of three in the old forest. In contrast to the small impact on C storage, afforestation strongly influenced the composition and quality of the soil organic matter (SOM). With increasing stand age, δ13C values of the SOM became consistently more positive, which can be interpreted as a gradual replacement of grass- by spruce-derived C. Fine roots of spruce were enriched in 13C, in lignin and had a higher C/N ratio in comparison to grass roots. As a consequence, SOM quality as indicated by the lower fraction of readily decomposable (labile) SOM and higher C:N ratios declined after the land-use change. Furthermore, spruce plantation induced a less favorable microclimate for microbial activity with the average soil temperature during the growing season being 5xa0°C lower in the spruce stands than in the pasture. In situ soil respiration was approximately 50xa0% lower after the land use conversion, which we primarily attribute to the colder conditions and the lower SOM quality, but also to drier soils (−25xa0%) and to a decreased fine root biomass (−40xa0%). In summary, afforestation on subalpine pastures only moderately affected SOC storage as compared to the large C sink in tree biomass. In contrast, SOC cycling rates strongly decreased as a result of a less favorable microclimate for decomposition of SOM, a lower C input by roots, and a lower litter quality.

Macronutrient inputs by litterfall as opposed to atmospheric deposition into two contrasting chestnut forest stands in southern Switzerland

Abstract The litterfall in two contrasting chestnut stands in Copera and Morbio, southern Switzerland, was collected between October 1994 and September 1995 and separated into the fractions leaves, branches, flowers, fruits and husks. The fractions were weighed and the concentrations of N, Ca, Mg, K, and C were measured. Wet deposition and throughfall of N, Ca, Mg, and K were measured and the corresponding dry deposition was estimated using multiple linear regression analysis. The total amount of litter in the two chestnut stands was 5.8 and 7.6xa0Mgxa0ha −1 xa0a −1 in Morbio and Copera, respectively. Leaves are the main constituents of the total litter. The proportions of the other litter fractions depend mainly on the former cultivation and management practice of the forests, the structure of the stand and the age of the trees. Regarding the concentration of the macronutrients in the litter fractions, leaves are richest in N and Ca, fruits in N and K. In husks, N dominates and Ca, and K do not differ very much. In branches, the N- and Ca-concentrations dominate by far, and the flowers are the richest in N. The return of macronutrients by litterfall is relevant: it is 6.6, 1.5, 0.6, and 1.3xa0kmolxa0ha −1 xa0a −1 for N, Ca, Mg, and K, respectively, in Copera. The corresponding values in Morbio are 3.8, 2.2, 0.5, and 0.3xa0kmolxa0ha −1 xa0a −1 . The return by litterfall is considerably higher than the import by atmospheric deposition (factors 3.8–5.1 in Morbio and Copera, respectively). In Copera, an amount of nutrient cations which corresponds to approximately 35% of the exchangeable stock in the soil is kept in the ecosystem internal cycle. This is highly relevant with respect to the buffering of acidity and for the prevention of nutrient cation loss due to acidification.

Aboveground vertebrate and invertebrate herbivore impact on net N mineralization in subalpine grasslands

Aboveground herbivores have strong effects on grassland nitrogen (N) cycling. They can accelerate or slow down soil net N mineralization depending on ecosystem productivity and grazing intensity. Yet, most studies only consider either ungulates or invertebrate herbivores, but not the combined effect of several functionally different vertebrate and invertebrate herbivore species or guilds. We assessed how a diverse herbivore community affects net N mineralization in subalpine grasslands. By using size-selective fences, we progressively excluded large, medium, and small mammals, as well as invertebrates from two vegetation types, and assessed how the exclosure types (ET) affected net N mineralization. The two vegetation types differed in long-term management (centuries), forage quality, and grazing history and intensity. To gain a more mechanistic understanding of how herbivores affect net N mineralization, we linked mineralization to soil abiotic (temperature; moisture; NO3-, NH4+, and total inorganic N concentrations/pools; C, N, P concentrations; pH; bulk density), soil biotic (microbial biomass; abundance of collembolans, mites, and nematodes) and plant (shoot and root biomass; consumption; plant C, N, and fiber content; plant N pool) properties. Net N mineralization differed between ET, but not between vegetation types. Thus, short-term changes in herbivore community composition and, therefore, in grazing intensity had a stronger effect on net N mineralization than long-term management and grazing history. We found highest N mineralization values when only invertebrates were present, suggesting that mammals had a negative effect on net N mineralization. Of the variables included in our analyses, only mite abundance and aboveground plant biomass explained variation in net N mineralization among ET. Abundances of both mites and leaf-sucking invertebrates were positively correlated with aboveground plant biomass, and biomass increased with progressive exclusion. The negative impact of mammals on net N mineralization may be related partially to (1) differences in the amount of plant material (litter) returned to the belowground subsystem, which induced a positive bottom-up effect on mite abundance, and (2) alterations in the amount and/or distribution of dung, urine, and food waste. Thus, our results clearly show that short-term alterations of the aboveground herbivore community can strongly impact nutrient cycling within ecosystems independent of long-term management and grazing history.

Does one model fit all? Patterns of beech mortality in natural forests of three European regions

Large uncertainties characterize forest development under global climate change. Although recent studies have found widespread increased tree mortality, the patterns and processes associated with tree death remain poorly understood, thus restricting accurate mortality predictions. Yet, projections of future forest dynamics depend critically on robust mortality models, preferably based on empirical data rather than theoretical, not well-constrained assumptions. We developed parsimonious mortality models for individual beech (Fagus sylvatica L.) trees and evaluated their potential for incorporation in dynamic vegetation models (DVMs). We used inventory data from nearly 19,000 trees from unmanaged forests in Switzerland, Germany, and Ukraine, representing the largest dataset used to date for calibrating such models. Tree death was modelled as a function of size and growth, i.e., stem diameter (dbh) and relative basal area increment (relBAI), using generalized logistic regression accounting for unequal re-measurement intervals. To explain the spatial and temporal variability in mortality patterns, we considered a large set of environmental and stand characteristics. Validation with independent datasets was performed to assess model generality. Our results demonstrate strong variability in beech mortality that was independent of environmental or stand characteristics. Mortality patterns in Swiss and German strict forest reserves were dominated by competition processes as indicated by J-shaped mortality over tree size and growth. The Ukrainian primeval beech forest was additionally characterized by windthrow and a U-shaped size-mortality function. Unlike the mortality model based on Ukrainian data, the Swiss and German models achieved good discrimination and acceptable transferability when validated against each other. We thus recommend these two models to be incorporated and examined in DVMs. Their mortality predictions respond to climate change via tree growth, which is sufficient to capture the adverse effects of water availability and competition on the mortality probability of beech under current conditions.