Dr. Mathias Mayer

Background and aims: Alpine shrublands are critical for global carbon dynamics due to their widespread occurrence in cold-climate regions and vulnerability to environmental shifts, including increased nitrogen deposition. Although nitrogen deposition affects litter turnover and accumulation, the precise mechanisms governing litter dynamics, including production, chemical composition, and decomposition rates, remain uncertain for these ecosystems.
Methods: To address this knowledge gap, our study investigated the effects of different nitrogen additions on litter production, chemistry, and decomposition rates in an alpine shrubland ecosystem on the eastern margin of the Qinghai–Tibetan Plateau of China over a four-year period.
Results: Our results showed that nitrogen addition did not significantly affect litter production or chemical properties, including carbon, nitrogen, phosphorus, lignin, and cellulose concentrations. Consequently, the annual input of litter-derived carbon and nutrients remained unchanged. However, we observed a significant reduction in litter decomposition rates at nitrogen additions of 50 and 100 kg ha⁻¹ yr⁻¹, whereas no such effect was observed at nitrogen additions of 20 kg ha⁻¹ yr⁻¹.
Conclusions: Our study revealed that high nitrogen deposition reduces litter decomposition in alpine shrublands, which coincides with increased litter accumulation, with consequences for carbon and nutrient cycling.

Treelines advance due to climate warming. The impacts of this vegetation shift on plant-soil nutrient cycling are still uncertain, yet highly relevant as nutrient availability stimulates tree growth. Here, we investigated nitrogen (N) and phosphorus (P) in plant and soil pools along two tundra-forest transects on Kola Peninsula, Russia, with a documented elevation shift of birch-dominated treeline by 70 m during the last 50 years. Results show that although total N and P stocks in the soil-plant system did not change with elevation, their distribution was significantly altered. With the transition from high-elevation tundra to low-elevation forest, P stocks in stones decreased, possibly reflecting enhanced weathering. In contrast, N and P stocks in plant biomass approximately tripled and available P and N in the soil increased fivefold toward the forest. This was paralleled by decreasing carbon (C)-to-nutrient ratios in foliage and litter, smaller C:N:P ratios in microbial biomass, and lower enzymatic activities related to N and P acquisition in forest soils. An incubation experiment further demonstrated manifold higher N and P net mineralization rates in litter and soil in forest compared to tundra, likely due to smaller C:N:P ratios in decomposing organic matter. Overall, our results show that forest expansion increases the mobilization of available nutrients through enhanced weathering and positive plant-soil feedback, with nutrient-rich forest litter releasing greater amounts of N and P upon decomposition. While the low N and P availability in tundra may retard treeline advances, its improvement toward the forest likely promotes tree growth and forest development.

Aim: Forest disturbances are increasing around the globe due to changes in climate and management, deteriorating forests' carbon sink strength. Estimates of global forest carbon budgets account for losses of plant biomass but often neglect the effects of disturbances on soil organic carbon (SOC). Here, we aimed to quantify and conceptualize SOC losses in response to different disturbance agents on a global scale.
Location: Global.
Time Period: 1983–2022.
Major Taxa Studied: Forest soils.
Methods: We conducted a comprehensive global analysis of the effects of harvesting, wildfires, windstorms and insect infestations on forest SOC stocks in the surface organic layer and top mineral soil, synthesizing 927 paired observations from 151 existing field studies worldwide. We further used global mapping to assess potential SOC losses upon disturbance. Results: We found that both natural and anthropogenic forest disturbances can cause large SOC losses up to 60 Mg ha⁻¹. On average, the largest SOC losses were found after wildfires, followed by disturbances from windstorms, harvests and insects. However, initial carbon stock size, rather than disturbance agent, had the strongest influence on the magnitude of SOC losses. SOC losses were greatest in cold-climate forests (boreal and mountainous regions) with large accumulations of organic matter on or near the soil surface. Negative effects are present for at least four decades post-disturbance. In contrast, forests with small initial SOC stocks experienced quantitatively lower carbon losses, and their stocks returned to pre-disturbance levels more quickly.
Main Conclusions: Our results indicate that the more carbon is stored in the forest's organic layers and top mineral soils, the more carbon will be lost after disturbance. Robust estimates of forest carbon budgets must therefore consider disturbance-induced SOC losses, which strongly depend on site-specific stocks. Particularly in cold-climate forests, these disturbance-related losses may challenge forest management efforts to sequester CO₂.

Fine root decomposition is an important driver of forest carbon (C) and nutrient cycling. Harvesting operations may affect fine root decomposition rates by altering root properties and environmental conditions, but our understanding of root dynamics is limited. In this study, we investigated the chemistry, mass loss, element release (C, nitrogen (N), and phosphorus (P)), and compound release (lignin and cellulose) of decaying fine roots in a 26 year-old Chinese fir plantation seven years after low- and high-intensity thinning (30% and 70% tree removal) using two root size classes (< 1 mm and 1–2 mm diameter). Low-intensity thinning (LIT) did not affect mass loss in either fine root class or the release of fine root elements or compounds during decomposition. Similarly, high-intensity thinning (HIT) had no effect on the decomposition of large fine roots. However, compared with LIT and no thinning, HIT reduced the decay rates and lignin and cellulose losses of small fine roots. This reduction was related to an increase in the root lignocellulose index (lignin/[lignin+cellulose]) and a decrease in soil invertase activity. Interestingly, thinning did not affect root C, N, or P loss during decomposition. In summary, our results suggest that thinning intensity as well as root size and chemistry should be considered when studying fine root dynamics in managed forests.

• Background and Aims Mixed forest plantations are increasingly recognized for their role in mitigating the impacts of climate change and enhancing ecosystem resilience. Yet, there remains a significant gap in understanding the early-stage dynamics of species trait diversity and interspecies interactions, particularly in pure deciduous mixtures. This study aims to explore the timing and mechanisms by which trait diversity of deciduous species and competitive interactions influence yield, carbon allocation and space occupation in mixed forests, both above and below ground.
• Methods A forest inventory was conducted in planted monocultures, two-species and four-species mixtures of European Acer, Tilia, Carpinus and Quercus, representing a spectrum from acquisitive to conservative tree species. Effects of competition were assessed with linear mixed-effects models at the level of biomass and space acquisition, including leaf, canopy, stem and fine root traits.
• Key Results Early above-ground growth effects were observed 6 years post-planting, with significant biomass accumulation after 8 years, strongly influenced by species composition. Mixtures, especially with acquisitive species, exhibited above-ground overyielding, 1.5–1.9 times higher than monocultures. Fine roots showed substantial overyielding in high-diversity stands. Biomass allocation was species specific and varied markedly by tree size and the level of diversity and between acquisitive Acer and the more conservative species. No root segregation was found.
• Conclusions Our findings underscore the crucial role of species trait diversity in enhancing productivity in mixed deciduous forest plantations. Allometric changes highlight the need to differentiate between (active) acclimatizations and (passive) tree size-related changes, but illustrate major consequences of competitive interactions for the functional relationship between leaves, stem and roots. This study points towards the significant contributions of both above- and below-ground components to overall productivity of planted mixed-species forests.

Understanding the effects of timber harvesting on soil respiration, including its autotrophic and heterotrophic components and their temperature sensitivity, is crucial for predicting how forest management affects the carbon cycle. Here, we conducted a meta-analysis to assess these effects on a global scale, synthesizing data from 1656 paired observations from 143 studies worldwide. On average, harvesting increased soil respiration by 6.0 %, most significantly in coniferous forests and subtropical regions. The response of total soil respiration was more closely coupled to changes in its heterotrophic than in its autotrophic component. The positive effects of harvesting on both respiration components decreased with increasing harvest intensity and were positively correlated with changes in soil nitrogen, root biomass, and microbial biomass carbon. Harvesting reduced the temperature sensitivity of soil respiration by 6.4 %, particularly in coniferous forests and temperate regions. The temperature sensitivity of soil autotrophic respiration increased in the first years after harvesting compared to the control but was significantly lower in later stages (c. > 6 years) after harvesting. Furthermore, the effects of harvesting on soil respiration, its components and temperature sensitivity varied greatly between post-harvest treatments and seasons of measurement. The results of our synthesis provide a basis for refining ecosystem models to better predict soil carbon dynamics in harvested forests on a global scale.

Storms represent a major disturbance factor in forest ecosystems, but the effects of windthrows on soil organic carbon (SOC) stocks are poorly quantified. Here, we assessed the SOC stocks of windthrown forests at 19 sites across Switzerland spanning an elevation gradient from 420 to 1550 m, encompassing a strong climatic gradient. Results show that the effect size of disturbance on SOC stocks increases with the size of the initial SOC stocks. The largest windthrow-induced SOC losses of up to 29 t C ha^-1 occurred in high-elevation forests with a harsh climate developing thick organic layers. In contrast, SOC stocks of low-elevation forests with thin organic layers were hardly affected. A mineralization study further revealed high elevation forests to store higher amounts of easily mineralizable C in thick organic layers that got lost following windthrow. These findings are supported by a meta-analysis of available windthrow studies, showing an increase of storm-induced SOC losses with the size of the initial SOC stocks. Modelling simulations further indicate longer-lasting SOC losses and a slower recovery of SOC stocks after windthrow at high compared to low elevations, due to a slower regeneration of mountain forests and associated lower C inputs into soils in a harsh climate. Upscaling the experimental findings/observed patterns by linking them to a data base of Swiss forest soils shows a total SOC loss of ∼0.4 Mt. C for the whole forested area of Switzerland after two major storm events, counteracting the forest net carbon sink of decades. Our study provides strong evidence that the vulnerability of SOC stocks to windthrow is particularly high in forests featuring thick and slowly forming organic layers, such as mountain soils. Thus, the risk of losing SOC to more frequent windthrows in mountain forests strongly limits their potential to mitigate climate change.

Ectomycorrhizal (ECM) fungi can both accelerate and decelerate decomposition of organic matter in forest soils, but a mechanistic understanding of this differential influence is limited.
Here, we tested how ECM fungi affect decomposition along a natural fertility gradient in a temperate forest of European beech. Trees were girdled to reduce belowground carbon supply to the soil.
Girdling shifted soil fungal community composition and decreased hyphal biomass production and soil CO₂ efflux, indicating a reduced ECM fungal activity. Girdling also affected decomposition processes, but the effects depended on fertility. Our results indicate that ECM fungi decelerate decomposition under conditions of low fertility while under conditions of high fertility ECM fungi and their host roots have an accelerating effect.
We conclude that both acceleration and deceleration of decomposition of organic matter by ECM fungi can occur within a forest, with soil fertility determining the direction and magnitude of these effects. We suggest a positive feedback between fertility, stand productivity and soil carbon and nitrogen dynamics that is mediated to a large extent by ECM fungi.

Forests in the Himalayas are a major carbon store, but are under threat due to changes in precipitation regime. To simulate a precipitation decline, throughfall-exclusion (TFE) shelters were applied during three consecutive monsoon seasons in an oak forest (2.650 m a.s.l.) and a conifer-dominated forest (3.260 m a.s.l.) in central Bhutan. Leaf water potentials, tree mortality, stem increment, soil CO₂ efflux, litterfall and fine root dynamics were assessed. TFE significantly and consistently decreased topsoil (0–30 cm) moisture and leaf water potentials of Quercus lanata and Quercus griffithii (lower elevation), and to a lesser extend those of Tsuga dumosa and Quercus semecarpifolia, (higher elevation). TFE did not impose tree mortality. Stem increment remained unaffected until the second TFE year, but showed reductions during the third year with Tsuga dumosa being most severely affected (-60%). Standing fine root biomass stocks were hardly affected by TFE. Increased root necromass and faster fine root growth in the lower elevation forest suggest that the oak trees increased C allocation below ground. Soil CO₂ efflux sharply declined during all three TFE years in both forests. Above ground litter input was unaffected by TFE until the second treatment year. Overall, both forest ecosystems appeared highly resistant to the imposed soil drying, with no signs of tree mortality and stable living root biomass stocks.

Der Boden ist eine zentrale Schnittstelle in Waldökosystemen. Er erfüllt wichtige Funktionen als Lebensraum, als Speicher für Kohlenstoff und Nährstoffe, und er reguliert die Wasser- und Stoffkreisläufe. Für alle Bodenfunktionen spielt die organische Bodensubstanz eine Schlüsselrolle. Aufgrund des feuchten und kühlen Klimas sowie der relativ nachhaltigen Waldnutzung speichern Schweizer Waldböden pro Flächeneinheit die grössten Vorräte organischer Bodensubstanz Europas. Dieser Vorrat hat sich über Tausende Jahre hinweg aufgebaut. Der Klimawandel - höhere Temperaturen und eine intensivere Trockenheit - sowie grossflächige Störungen, beispielsweise durch Windwürfe und Kahlschläge, gefährden die organische Bodensubstanz. Eine bodenschonende Waldbewirtschaftung sowie eine permanente Bestockung mit standortsangepassten Baumarten tragen zum Erhalt dieser wichtigen Ressource bei.

Soil represents an essential component in forest ecosystems. It fulfills key functions as living space, storage for carbon and nutrients, regulation of water and element cycles. Soil organic matter plays a key role for all soil functions. Swiss forest soils contain the greatest organic matter stocks per unit area of all European countries due to the relatively cool and humid climate and the overall sustainable forest management. The stocks have built up over thousands of years, but are put at risk by climate change - higher temperatures and more intense drought - as well as disturbances by windthrows or clear cuts. A soil-friendly forest harvest, a continuous forest cover, as well as the promotion of a structure- and species-rich forest are all contributing to preserve soil organic matter as an essential resource in the soil.

Wälder sind zunehmend von Sturmschäden betroffen. In einem Projekt der Eidgenössischen Forschungsanstalt WSL wurde untersucht, wie sich Windwürfe auf die Kohlenstoffspeicherung im Waldboden auswirken. Hoch gelegene Nadelwälder sind besonders anfällig für Kohlenstoffverluste aus dem Boden. Im Unterschied zu Wäldern tieferer Lagen werden dort grössere Mengen an Kohlenstoff in rein organischen, mächtigen Humusauflagen gespeichert. Eine Stabilisierung durch mineralische Interaktionen fehlt weitgehend, was die Humusauflagen empfindlich macht für Kohlenstoffverluste durch störungsbedingte Veränderungen des Bestandesklimas. Die langsame Baumverjüngung und die geringen Streueinträge in den Boden bremsen den Humusaufbau nach einer Störung zusätzlich. Eine weitere Zunahme von Windwürfen könnte die Bodenkohlenstoffvorräte in Bergwäldern drastisch reduzieren, was negative Auswirkungen auf das Klima hätte. Darüber hinaus könnten störungsbedingte Humusverluste die Bodenqualität nachhaltig verschlechtern.

Storms represent a major disturbance factor in forest ecosystems, but the effects of windthrows on soil organic carbon stocks are poorly known. The Swiss Federal Institute WSL assessed the soil organic carbon stocks of windthrown forests and neighbouring unaffected stands across Switzerland. The largest soil carbon losses occurred in high-elevation forests with thick organic layers. In contrast, soil organic carbon stocks of low-elevation forests with thin organic layers were hardly affected. The likely reason for this pattern is the high amount of easily mineralizable carbon in thick organic layers which got lost following windthrow. At low elevations, on the other hand, a greater carbon fraction may be stabilized by mineral interactions preventing fast decomposition. Modelling simulations further show longer-lasting soil carbon losses and a slower recovery of carbon stocks after windthrow at high elevations compared to low elevations, due to a slower regeneration of mountain forests and associated lower carbon inputs into soils. The study provides strong empirical evidence that the vulnerability of soil organic carbon stocks to windthrow is particularly high in mountain forests featuring thick and slowly forming organic layers.

Temperate forests are increasingly subject to natural disturbance by stand replacing windthrows or bark-beetle attacks. Forests are commonly salvage logged after disturbance, whereby substantial parts of biological legacies, such as surviving trees and deadwood, are removed. Despite increasing concerns about the ecological consequences of salvage logging operations, our knowledge on the effects on the soil microbiome and associated functioning remains limited. Here, we studied soil fungal communities, decomposition processes, and soil organic matter dynamics in 21 intact or disturbed, temperate Norway spruce stands about one decade after they were damaged by windthrow or bark-beetle attacks. Disturbed stands comprised different post-disturbance management, i.e. deadwood retention and salvage logged plots. We used high-throughput sequencing and ergosterol measurements to explore fungal communities and biomass, and enzyme assays to study decomposition processes. Disturbance shifted soil fungal communities from ectomycorrhizal to saprotrophic dominated assemblages. Fungal biomass declined with decreasing tree abundance after disturbance. Activities of organic matter degrading enzymes declined by ca. 30–80% after disturbance. The relative abundance of ectomycorrhizal fungi was positively related to enzymatic activities. Tree biomass parameters and amounts of deadwood retained were positively related to fungal biomass, certain ectomycorrhizal taxa, and relative ectomycorrhizal fungal abundance among disturbed stands, which, in turn, was associated with higher enzymatic activities. Our findings demonstrate a significant response of soil fungal communities to natural forest disturbance and salvage logging, with consequences for decomposition and soil organic matter dynamics. We conclude that the retention of surviving trees and deadwood as biological legacies attenuated associated changes to a significant extent, highlighting their importance for the preservation of ectomycorrhizal fungi and the maintenance of decomposition processes after disturbance.

Forests on steep slopes constitute a significant proportion of European mountain areas and are important as production and protection forests. This study describes the soil fungal community structure in a European beech-dominated mountain forest stands in the Northern Calcareous Alps and investigates how it is determined by season and soil properties. Samples were collected at high spatial resolution in an area of ca. 100 m × 700 m in May (spring) and August (summer). Illumina MiSeq high-throughput sequencing of the ITS2-region revealed distinct patterns for the soil fungal communities. In contrast to other studies from temperate European beech forest stands, Ascomycota dominated the highly diverse fungal community, while ectomycorrhizal fungi were of lower abundance. Russulaceae, which are often among the dominant ectomycorrhizal fungi associated with European beech, were absent from all samples. Potentially plant pathogenic fungi were more prevalent than previously reported. Only subtle seasonal differences were found between fungal communities in spring and summer. Especially, dominant saprotrophic taxa were largely unaffected by season, while slightly stronger effects were observed for ectomycorrhizal fungi. Soil characteristics like pH and organic carbon content, on the other hand, strongly shaped abundant taxa among the saprotrophic fungal community.

Fungi are known to exert a significant influence over soil organic matter (SOM) turnover, however understanding of the effects of fungal community structure on SOM dynamics and its consequences for ecosystem fertility is fragmentary.
Here we studied soil fungal guilds and SOM decomposition processes along a fertility gradient in a temperate mountain beech forest. High-throughput sequencing was used to investigate fungal communities. Carbon and nitrogen stocks, enzymatic activity and microbial respiration were measured.
While ectomycorrhizal fungal abundance was not related to fertility, saprotrophic ascomycetes showed higher relative abundances under more fertile conditions. The activity of oxidising enzymes and respiration rates in mineral soil were related positively to fertility and saprotrophic fungi. In addition, organic layer carbon and nitrogen stocks were lower on the more fertile plots, although tree biomass and litter input were higher. Together, the results indicated a faster SOM turnover at the fertile end of the gradient.
We suggest that there is a positive feedback mechanism between SOM turnover and fertility that is mediated by soil fungi to a significant extent. By underlining the importance of fungi for soil fertility and plant growth, these findings furthermore emphasise the dependency of carbon cycling on fungal communities below ground.

The relatively poor simulation of the below-ground processes is a severe drawback for many ecosystem models, especially when predicting responses to climate change and management. For a meaningful estimation of ecosystem production and the cycling of water, energy, nutrients and carbon, the integration of soil processes and the exchanges at the surface is crucial. It is increasingly recognized that soil biota play an important role in soil organic carbon and nutrient cycling, shaping soil structure and hydrological properties through their activity, and in water and nutrient uptake by plants through mycorrhizal processes. In this article, we review the main soil biological actors (microbiota, fauna and roots) and their effects on soil functioning. We review to what extent they have been included in soil models and propose which of them could be included in ecosystem models. We show that the model representation of the soil food web, the impact of soil ecosystem engineers on soil structure and the related effects on hydrology and soil organic matter (SOM) stabilization are key issues in improving ecosystem-scale soil representation in models. Finally, we describe a new core model concept (KEYLINK) that integrates insights from SOM models, structural models and food web models to simulate the living soil at an ecosystem scale.

Belowground competition is an important structuring force in terrestrial plant communities. Uncertainties remain about the plasticity of functional root traits under competition, especially comparing interspecific vs. intraspecific situations. This study addresses the plasticity of fine root traits of competing Acer pseudoplatanus L. and Fagus sylvatica L. seedlings in nutrient-rich soil patches. Seedlings’ roots were grown in a competition chamber experiment in which root growth (biomass), morphological and architectural fine roots traits, and potential activities of four extracellular enzymes were analyzed. Competition chambers with one, two conspecific, or two allospecific roots were established, and fertilized to create a nutrient 'hotspot'. Interspecific competition significantly reduced fine root growth in Fagus only, while intraspecific competition had no significant effect on the fine root biomass of either species. Competition reduced root nitrogen concentration and specific root respiration of both species. Potential extracellular enzymatic activities of β-glucosidase (BG) and N-acetyl-glucosaminidase (NAG) were lower in ectomycorrhizal Fagus roots competing with Acer. Acer fine roots had greater diameter and tip densities under intraspecific competition. Fagus root traits were generally more plastic than those of Acer, but no differences in trait plasticity were found between competitive situations. Compared to Acer, Fagus roots possessed a greater plasticity of all studied traits but coarse root biomass. However, this high plasticity did not result in directed trait value changes under interspecific competition, but Fagus roots grew less and realized lower N concentrations in comparison to competing Acer roots. The plasticity of root traits of both species was thus found to be highly species- but not competitor-specific. By showing that both con- and allospecific roots had similar effects on target root growth and most trait values, our data sheds light on the paradigm that the intensity of intraspecific competition is greater than those of interspecific competition belowground.

Almost half of the total organic carbon (C) in terrestrial ecosystems is stored in forest soils. By altering rates of input or release of C from soils, forest management activities can influence soil C stocks in forests. In this review, we synthesize current evidence regarding the influences of 13 common forest management practices on forest soil C stocks. Afforestation of former croplands generally increases soil C stocks, whereas on former grasslands and peatlands, soil C stocks are unchanged or even reduced following afforestation. The conversion of primary forests to secondary forests generally reduces soil C stocks, particularly if the land is converted to an agricultural land-use prior to reforestation. Harvesting, particularly clear-cut harvesting, generally results in a reduction in soil C stocks, particularly in the forest floor and upper mineral soil. Removal of residues by harvesting whole-trees and stumps negatively affects soil C stocks. Soil disturbance from site preparation decreases soil C stocks, particularly in the organic top soil, however improved growth of tree seedlings may outweigh soil C losses over a rotation. Nitrogen (N) addition has an overall positive effect on soil C stocks across a wide range of forest ecosystems. Likewise, higher stocks and faster accumulation of soil C occur under tree species with N-fixing associates. Stocks and accumulation rates of soil C also differ under different tree species, with coniferous species accumulating more C in the forest floor and broadleaved species tending to store more C in the mineral soil. There is some evidence that increased tree species diversity could positively affect soil C stocks in temperate and subtropical forests, but tree species identity, particularly N-fixing species, seems to have a stronger impact on soil C stocks than tree species diversity. Management of stand density and thinning have small effects on forest soil C stocks. In forests with high populations of ungulate herbivores, reduction in herbivory levels can increase soil C stocks. Removal of plant biomass for fodder and fuel is related to a reduction in the soil C stocks. Fire management practices such as prescribed burning reduce soil C stocks, but less so than wildfires which are more intense. For each practice, we identify existing gaps in knowledge and suggest research to address the gaps.