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Global environmental changes and the subsequent biodiversity loss has raised concerns over the consequences for the functioning of ecosystems and human well-being. This thesis provides new mechanistic insights into the role of tree diversity in regulating forest productivity and forests’ responses to climate change. The thesis also addresses the overlooked functional role of ecological continuity in mediating ecosystem processes in the context of multiple global environmental changes. The findings of the thesis emphasize the need to retain the functional integrity of forest ecosystem by preserving biodiversity and acknowledging the ecological memory forests.
Gegenwärtige atmogene N-Einträge in die Lüneburger Heide überschreiten die durch den critical-load für Sandheiden definierte Belastungsgrenze von 10-20 kg ha-1 und zeigen den Handlungsbedarf für eine langfristige Erhaltung dieser geschützten Lebensräume. Die traditionelle Schafbeweidung im NSG Lüneburger Heide kann diese hohen atmogenen N-Einträge kompensieren. Das Herdenmanagement minimiert den Nährstoffeintrag über die Exkremente der Tiere, so dass diesem Transfer nur eine geringe Bedeutung zukommt. Die N-Austräge über das Sickerwasser vor, während und nach den Pflegemaßnahmen spielen nur eine untergeordnete Rolle. Extensive Pflegemaßnahmen wie die Mahd und das kontrollierte Brennen sind nicht geeignet die atmogenen N-Einträge langfristig zu kompensieren. Intensiven Pflegemaßnahmen (Schoppern, Plaggen) gelingt dieses, benötigen aber einen zu langen Bearbeitungszyklus, innerhalb dessen kombinierte Verjüngungsmaßnahmen notwenig werden. Die Pflegemaßnahmen haben Auswirkungen auf die N- und P-Vorräte im Heidesystem und führen großteils zu einer P-Verarmung. Das kontrollierte Brennen beeinflusst den P-Haushalt am geringsten. Eine Kombination des Feuereinsatzes mit anderen extensiven Maßnahmen erscheint daher sinnvoll.
Assessment of forest functionality and the effectiveness of forest management and certification
(2021)
Forest ecosystems are complex systems that develop inherent structures and processes relevant for their functioning and the provisioning of ecosystem services that contribute to human wellbeing. So far, forest management focused on timber production while other services were less rewarded. With increasing climate change impacts, especially regulating ecosystem services such as microclimate regulation are ever more relevant to maintain forest functions and services. A key question is how forest management supports or undermines the ecosystems’ capacity to maintain those functions and services. Forest management implies silvicultural interventions such as thinning and timber harvesting and ranges from single tree extraction to large clearcuts as well as forest reserves without active forest operations and shape the character of forest ecosystems (e.g. natural versus planted forests). Artificial plantings, monocultures and management for economic timber production simplify forest structures and impair ecosystem resilience, resistance and the existence of forests but also the services essential for the prosperity and health of humanity. Efforts to reduce the negative impacts and attempts to safeguard forest functions are manifold and include compulsory national and international guidelines and regulations for forest management, conventions, but also voluntary mechanisms such as certification systems.
The main objective of this thesis was the development of a concept to assess the functionality of forests and to evaluate the effectiveness of forest ecosystem management including certification. An ecosystem-based and participatory methodology, named ECOSEFFECT, was developed. The method comprises a theoretical and an empirical plausibility analysis. It was applied to the Russian National FSC Standard in the Arkhangelsk Region of the Russian Federation – where boreal forests are exploited to meet Europe's demand for timber. In addition, the influence of forestry interventions on temperature regulation in Scots pine and European beech forests in Germany was assessed during two extreme hot and dry years in 2018 and 2019.
Microclimate regulation is a suitable proxy for forest functionality and can be applied easily to evaluate the effectiveness of forest management in safeguarding regulating forest functions relevant under climate change. Microclimate represents the most decisive factor differentiating clearcuts and primary forests. Thus, the assessment of forest microclimate regulation serves as convenient tool to illustrate forest functionality. In the boreal and temperate forests studied in the frame of this thesis, timber harvesting reduced the capacity to self-regulate forests’ microclimate and thus impair a crucial part of ecosystem functionality. Changes in structural forest characteristics influenced by forest management and silviculture significantly affect microclimatic conditions and therefore forest ecosystems’ vulnerability to climate change. Canopy coverage and the number of cut trees were most relevant for cooling maximum summer temperature in pine and beech forests in northern Germany. Maximum temperature measured at ground level increased by 0.21 – 0.34 K when 100 trees were cut. Opening the forest canopy by 10 % caused an increase of maximum temperature at ground-level by 0.53 K (including pine and beech stands). Relative temperature cooling capacity decreased with increasing wood harvest activities and dropped below average values when more than 656 trees per hectare (in 2018; and 867 trees in 2019) were felled. In pine stands with a canopy cover below 82 % the relative temperature buffering capacity was lower than the average. Mean maximum temperature measured at ground-level and in 1.3 m was highest in a pine-dominated sample plots with relatively low stand volume (177 m3 ha-1) and 9 K lower in a sample plot with relatively high stock volumes of F. sylvatica (> 565 m3 ha-1). During the hottest day in 2019, the difference in temperature peaks was more than 13 K for pine-dominated sample plots with relatively dense (72 %) and low (46 %) canopy cover.
The Russian FSC standard has the potential to improve forest management and ecological outcomes, but there are shortcomings in the precision of targeting actual problems and ecological commitment. In theory, FSC would transform forest management practices and induce positive changes and effective outcomes by addressing 75 % of the identified contributing factors including highly relevant factors and threats including large-scale (temporary) tree cover loss, which contributes to reducing about half of the identified stresses in the ecosystem. It is theoretically plausible that FSC prevents logging in high conservation value forests and intact forest landscapes, reduces the size and number of clearcuts, and prevents hydrological changes in the landscape. However, the standard was not sufficiently explicit and compulsory to generate a strong and positive influence on the identified problems and their drivers. Moreover, spatial data revealed, that the typical regular clearcut patterns of conventional timber harvesting continue to progress into the FSC-certified boreal forests, also if declared as ‘Intact Forest Landscape’. This results in the need to verify the assumptions and postulates on the ground as it remains unclear and questionable if functions and services of boreal forests are maintained when FSC-certified clearcutting continues. On the clearcuts, maximum temperature exceeded 36 °C and stayed below 30 °C in the closed primary forest. The number of days with temperatures above 25 °C at least doubled on clearcuts. Temperature cooling capacity was reduced by up to 14 % and temperature buffering capacity up to 60 %. The main reason why FSC-certified clearcuts do not differ from conventional clearcuts is that about 97 % of trees within equally large clearcut sites of up to 50 ha were removed. The spatial design of clearcuts, their size and the intensity of clearing as well as the density of skidding trails for timber extraction was not positively influenced by FSC-certification. Annual tree cover loss was lowest in non-certified areas. This means, that FSC may even contribute to an increased biomass removal within the clearcuts, which compromises the ecosystems’ capacity to recover and maintain ecological functions and services. The analysis of satellite-based data on tree cover loss showed that clearcutting causes secondary dieback in the surrounding of the cleared area. FSC-certification does not prevent the various negative impacts of clearcutting and thus fails to safeguard ecosystem functions. The postulated success in reducing identified environmental threats and stresses, e. g. through a smaller size of clearcuts, could not be verified on site. The empirical assessment does not support the hypothesis of effective improvements in the ecosystem. In practice, FSC-certification did not contribute to change clearcutting practices sufficiently to effectively improve the ecological performance. Sustainability standards that are unable to translate principles into effective outcomes fail in meeting the intended objectives of safeguarding ecosystem functioning. Clearcuts that carry sustainability labels are ecologically problematic and ineffective for the intended purpose of ecological sustainability.
The overexploitation of provisioning services, i.e. timber extraction, diminishes the ecosystems’ capacity to maintain other services of global significance. It also impairs ecosystem functions relevant to cope with and adapt to other stresses and disturbances that are rapidly increasing under climate change.
Forest management under climate change needs to apply precautionary principles and reduce further ecological risks such as secondary dieback and deterioration of regulating services that are relevant for the functioning of forests. Forest managers have to avoid ecological disimprovements by applying strict ecological principles with effective outcomes in order to maintain functional forests that regulate their own microclimate also as a basis for sustainable economic benefits.
The dissertation deals with the impact of nitrogen deposition on the functioning of heathland ecosystems. Special interests were the displacement of heather (Calluna vulgaris) by the purple moor-grass (Molinia caerulea) as well as the fate of nitrogen loads in dry heathland ecosystems. The results of the studies undertaken in the field and in the greenhouse are presented as five individual journal articles. The nature of nutrient limitation was studied by means of fertilisation experiments with nitrogen (N) and phosphorus for heather and purple moor-grass (Articles I and II). The impact of nitrogen deposition on the outcome of competition between these two species was analysed during a competition experiment in the greenhouse (Article III). The aim of a 15N tracer experiment was to determine the fate of nitrogen deposition as well as allocation patterns (Article IV). In addition, the response of purple moor-grass to the combined effects of nitrogen deposition and summer droughts was investigated in a second greenhouse experiment (Article V). The fertilisation experiments showed that the growth of heather as well as of purple moor-grass is predominantly limited by N (Articles I and II). However, the results of the competition experiment demonstrated that only purple moor-grass has the ability to benefit from additional N loads, which in turn gives the grass the opportunity to displace heather (Article III). Drought treatment resulted in strikingly reduced biomass production of purple moor-grass in N-fertilised pots, mainly as a result of dying aboveground biomass during dry periods (Article V). This striking susceptibility of purple moor-grass to the combination of nitrogen deposition and drought must be taken into account, when predicting future developments of dry heathlands. The results of the 15N tracer experiment showed that the investigated heath is still in an early stage of N saturation, as indicated by a high immobilisation capacity and negligible leaching losses of 15N (Article IV). The findings of the dissertation contribute to a better understanding of the processes underlying the encroachment of purple moor-grass in dry heathlands and can enhance heathland management. The results can also be used to to evaluate the current and future status of this ecosystem particularly with regard to the various stages of N saturation as well as in the determination of “Critical Loads”.
Restoration and management of abandoned, dry continental heathland and sandy grassland communities
(2018)
Land-use changes and long-term abandonment are main drivers that change ecosystem functioning and cause biodiversity loss of many semi-natural habitats, such as heathlands and grasslands. Traditional management measures such as low-intensity grazing created these semi-open landscapes and maintained a high species richness. However, from the middle of the 19th century onwards, traditional management practices began to decline due to socio-economic changes, and large areas of heathlands and grasslands were subjected to succession and reverted to woodland. Nowadays, dry heathlands as well as dry sandy grasslands are recognized as being of high conservation value and classified as ´habitats of community interest´ (´European dry heaths´, habitat code 4030; ´Xeric sand calcareous grasslands´, habitat code *6120). Whereas heathlands in the Atlantic biogeographical region have been in the centre of interest, dry heathland communities in the Continental biogeographical region have been widely neglected, even though they comprise 30 percent of all dry heaths as well as 89 percent of all dry sandy grasslands, respectively, in Europe. Thereby, the conservation status of both habitat types is listed as unfavourable-bad across the Continental biogeographical region. Surprisingly, no detailed studies are available on cost-efficient and sustainable restoration and management schemes to successfully restore and maintain highly degraded, long-abandoned Continental heaths and sandy grasslands, and thus, to counteract the poor conservation status of the habitat types. This shows the great need for research for the Continental biogeographical region (chapter I). Thus, the present thesis provides substantial knowledge about the population dynamics of the key plant species of dry heaths Calluna vulgaris (L.) HULL (henceforth referred to as Calluna) by investigating key processes in the biology of the species as well as about the restoration and management of long-abandoned, dry Continental heathland and sandy grassland communities. In order to better understand the process of successful Continental heathland restoration, I analysed the reproductive potential (seed production, soil seed bank, and germination ability of seeds) of degenerate Calluna stands as well as the effects of single and combined management options on the generative rejuvenation (i.e., recruitment and survival) of Calluna (chapter II). The results are based on a comprehensive three-year field experiment including the management options year-round, low-intensity cattle and horse grazing, one-time mowing and one-time shallow soil disturbances combined with greenhouse investigations on the soil seed bank content and germination ability of Calluna seeds. The results showed that even after long-term abandonment, seed production of degenerate Calluna stands and the germination ability of seeds proved to be high, being similar to Atlantic heathlands, whereas the soil seed bank is considerably reduced probably due to the dry conditions in the Continental region. In addition, low-intensity grazing with free-ranging robust breeds and the combination with one-time mowing at the beginning of the restoration process is an effective means of supporting the generative rejuvenation of this key plant species in degraded Continental heaths. The second study of this thesis (chapter III) focussed on the first-year establishment of Calluna in managed and unmanaged dry heaths and heaths in mosaics with dry sandy grasslands. The germination ability of seeds of different life-history phases of Calluna was analysed to determine if the predominance of the late life-history phase restricts the rejuvenation process of this key plant species. In addition, beside effects of management measures (year-round, low-intensity grazing, one-time mowing, one-time shallow soil disturbances) I analysed the most important safe site conditions that possibly influenced the germination and the first-year survival of Calluna. The results of the study combine field experiments with growth chamber investigations. I found that life-history phase of Calluna did not significantly affect seed germination and thus, the predominance of the degenerate life-history phase does not restrict the rejuvenation process. In addition, the results of my study revealed that grazing and thus trampling intensity must be temporarily and locally enhanced at the beginning of the restoration process of highly degraded heaths to increase safe site availability for successful Calluna establishment. Thereby, shadowing is the most important safe site condition that limit Calluna recruitment and survival in the first year in both degraded heaths as well as in mosaics with sandy grasslands, since seedlings receiving full sunlight die significantly more frequently than slightly or fully shaded seedlings. In the third study (chapter IV), I investigated the impacts of year-round low-intensity cattle and horse grazing on the development of the highly competitive grass Calamagrostis epigejos (henceforth referred to as Calamagrostis), as well as the vegetation structure and plant species richness of long-abandoned but nutrient-poor dry heathland and sandy grassland communities, their mosaics and Calamagrostis stands. Finally, I assessed the local conservation status of the habitat types after seven years of grazing in comparison to long-abandoned sites. The results are based on a comprehensive field study on two spatial scales (plot-level: 25 square meters, macroplot-level: 1 ha). I found that grazing successfully reduced the coverage and prevented the further spread of Calamagrostis, while simultaneously maintained or improved characteristic species richness and vegetation structure across the different nutrient-poor vegetation types over time, and thus enhanced the local conservation status of habitat types of community interest. In conclusion, the results of my studies considerably improved the understanding of dry, Continental heathland and sandy grassland restoration and management. They provide evidence that even after long-term management abandonment, year-round low-intensity cattle and horse grazing is a suitable management tool for restoring, maintaining and even improving nutrient-poor heathland and sandy grassland communities. However, at the beginning of the restoration process, additional management measures are necessary to faster restore abandoned habitats, especially highly degraded heaths.
Biodiversity loss could jeopardize ecosystem functioning. Yet, the evidences that support this demonstration have been mostly obtained in aquatic and grassland ecosystems. Howbiodiversity affects ecosystem functioning still remain largely unanswered in forests, particularly in subtropical broad-leaved evergreen forests (EBLF). Tree productivity, among a wealth of forest ecosystem functioning, is of particular interest because it reflects the carbon sink capacity and wood productivity. Biodiversity-productivity relationships have been usually investigated at community level. However, tree-tree interactions occur at small scale. Thus, local neighborhood approach may allow a better understanding of tree-tree interactions and their contributions to the effects of biodiversity on tree productivity / growth rates. This thesis aims to analyze the effects of biodiversity and the abiotic environmental factors on the tree growth rates using both local neighborhood and community-based approaches. Furthermore, tree growth rates vary among different tree species. Functional traits have been related to the species-specific growth rates to understand the effects of species identity. Therefore, I also evaluated the crown- and leaf traits to predict the interspecific difference in growth rates. For a better understanding of the mechanisms that underline the relationships of biodiversity and tree growth rates, data of high solution and along time series is required to scrutinize the tree-tree interactions. Thereupon, I evaluated the applicability of terrestrial laser scanning (TLS) in assessing the tree dendrometrics. This thesis was conducted in the Biodiversity Ecosystem Functioning (BEF)–China experiment, which is located in a mountainous subtropical region in southeast China. A total of 40 native broad-leaved tree species were planted. In the first study, I used the local neighborhood approach to analyze how local abiotic conditions (i.e. topographic and edaphic conditions) and local neighborhood (i.e. species diversity and competition by neighborhood) affect the annual growth rates of 6723 individual trees. The second study used the community approach to partition the effects of environmental factors (i.e. topographic and edaphic), functional diversity according to Rao’s quadratic entropy (FDQ) and community weight mean (CWM) of 41 functional traits on community tree growth rates. The main question of the third study was how the species-specific growth rates are related to five crown- and 12 leaf traits.
In the fourth study, I investigated 438 tree individuals for the congruence between the conventional direct field measurements and TLS measurements. It was found that tree growth rates were strongly influenced by the local topographic and edaphic conditions but not affected by the diversity of local neighborhood. In contrast, results obtained by using the community-based approach showed that FDQ and CWMs of various leaf traits rather than abiotic environmental factors had significant impact on the community means of growth rates. Tree-tree interactions already occur in early life stages of trees, which were evidenced by the significant effect of competition by local neighborhood. These findings imply that the effects of abiotic environmental factors may be more evident at local scale and biodiversity effects may vary at different spatial scales. The species-specific growth rates were found to be related to specific leaf traits but not to crown traits and were best explained by both types of traits in combination. This finding supports the niche theory and provides the evidence for using functional diversity to examine the BEF relationships. The TLS-retrieved total tree height, stem diameter at 5 cm above ground, and length and height of the longest branch were highly congruent with those obtained from direct measurements. It indicates that TLS is a promising tool for high resolution, non-destructive analyses of tree structures in young tree plantations. Being one of very few studies to incorporate the individual tree scale in examining the biodiversity-productivity relationships within the BEF researches, this thesis stresses the importance of using individual-tree based approach, functional diversity and TLS to find the evidences of explanatory mechanisms of the observed biodiversity and ecosystem functioning (e.g. tree growth rates) relationships. Biodiversity effects may evolve along the successional stages. Therefore, incorporating the interaction between biodiversity and time in analyzing BEF relationship is also encouraged.
Excessive fertilizer use leads to nutrient imbalances and losses of these to the environment through leaching, runoff and gaseous emissions. Nutrient use efficiency (NUE) in agriculture is often low and improving it could increase the sustainability of agricultural systems. The main aims of this thesis were to gain a better understanding of plant-soil-microbe interactions in order to improve agricultural NUEs. The studies included experimentally tested how crops respond to addition of high carbon amendments, fertilizer application rates and timing, and crop rotations. Furthermore, methods for measurement of roots were compared and a protocol for measurement of roots was developed.
The first experiment simulated an agricultural field using mesocosms. In this setting, we tested the effect of 4 previous crops (precrops), which either had or did not have a symbiosis with arbuscular mycorrhizal fungi (AMF)/rhizobia, on the focal crop (winter barley). We also tested the addition of high carbon amendments (wheat straw/sawdust) for immobilization of residual soil nitrogen (N) at harvest of the previous crop. Overall, the findings were that non-AMF precrops had a positive effect on winter barley yield compared to AMF precrops. Wheat straw reduced N leaching, whereas sawdust addition had a negative effect on the yield of winter barley.
Root traits are often measured in static environments, whereas agricultural fertilizer is applied once or multiple times at different crops growth stages. The second experiment tested the effect of different fertilizer (N/phosphorus (P)) application timings on plant traits grown in rhizoboxes. Overall, delaying N application had a more detrimental effect on plant biomass than delaying P application. The root system increased its root length initially due to N-deficiency, but was quickly thus N-limited that root length was relatively lower than the control group. This study emphasizes the need to dynamically measure roots for a mechanistic understanding of root responses to nutrients.
Because of the many root related measurements in the second experiment, a step-by-step method for measuring root traits under controlled and field conditions was developed and included in this thesis. This method paper describes precisely how root traits of interest can be measured, and helps with deciding which approach should be taken depending on the experimental design. Additionally, we compared the bias and accuracy of several popular root measurement methods. Although methods well correlated with a reference method, most methods tended to underestimate the total root length.
Overall, these results highlight the importance of crop choice in crop rotations and the plasticity of root systems in relation to nutrient application. Our results show high carbon amendments could reduce nitrate leaching after the harvest of crops, especially those with high risk of nitrate leaching, although they had only small impacts on yield. Future research should investigate the applicability in a farm setting, also taking into account financial and practical aspects. Non-AMF precrops could possibly increase yield of the next crop due to a shift to parasitism in agricultural fields, but whether this plays a large role in crop yields should be further investigated for specific soil, crop, and climate conditions. Our results also show the plasticity of root systems in response to nutrients. Understanding and using this plasticity can be useful for improving NUEs by optimizing fertilizer application and selecting root traits that are beneficial for specific environmental conditions.
European species-rich grasslands, which provide many ecosystem functions and services, are threatened both by land use intensification as well as land abandonment. The studies shown in this thesis tested the possible use of ecological knowledge to ensure hay productivity whilst maintaining diversity of grasslands, with a view to informing ecological restoration. The overall approach was to understand interactions between plants, to study diversity effects on productivity, and mainly investigate how plant functional groups that arrive first in the system can create priority effects that influence community productivity both above- and belowground. A grassland field experiment was established and monitored for four years, in order to verify the effects of manipulating the order of arrival of different plant functional groups, as well as the sown diversity level on productivity and methane yield. The overall findings were: a) sowing legumes first created priority effects aboveground (higher biomass) and belowground (lower root length), plants invested less in roots and more in shoots, b) priority effects were more consistent below than aboveground, c) sown diversity did not affect aboveground biomass, d) the order of arrival treatments indirectly affected methane yield by affecting the relative dominance of plant functional groups. Since we lack information on how legumes and non-legumes interact spatially belowground, (particularly related to root foraging) a controlled experiment was performed, using two grass species and one legume. The identity and location of the neighbours played a role in interactions, and the order plants arrived modulated it. When the focal species (grass) was growing with a legume it generally equated to the same outcome as not having a neighbour. Roots from the focal species grew more toward the legume than the grass neighbour, indicating a spatial component of facilitation. Since these studies involved root measurements, a method study was also conducted to verify how comparable and accurate are root length estimates obtained from different techniques. Results showed that the use of different methods can lead to different results, the studied methods did not have the same accuracy, and the automated methods can underestimate the root length. Overall, the results allow to conclude that different groups of plants arriving before others affected above and belowground biomass, roots may be key drivers during the creation of these priority effects, and interaction outcomes between plants depended on neighbour identity and location, modulated by the order they arrive in. Our results suggest that we can use priority effects by sowing different species or plant functional groups at different time to steer a community to a desired trajectory depending on the restoration goal (such as increasing biomass whilst maintaining diversity). However, there is a need to test contingency, potential, and long term impacts of such possible tools for restoration.