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A solid knowledge about nature is essential to understand the consequences of biodiversity loss, the limitation of natural resources and the need for a sustainable development. Inspired by these challenges, the researcher investigated in her dissertation seed predation, an important ecosystem function, as part of citizen science project. As seed predation has only rarely been investigated along urban-rural gradients and to integrate the question if the background (urban vs. rural) of primary school children affects their environmental knowledge, she selected study sites in and around Lüneburg and Hamburg, in Northern Germany. In her ecological experiments, it was found that slugs are important seed predators that independently of urbanization predated about 30% of all seeds in the anthropogenically used landscapes investigated. Also, for the first time, primary school children could be integrated in a citizen science approach into this research and it could be shown that even seven year old children can record data as reliable as a scientist. Finally, the researcher investigated the native species knowledge from the children taking part as citizen scientists in her research, considering possible differences due to their urban or rural background. Contrary to her expectation, the urban or rural background had no significant effect on the species knowledge. However, the work provides a good foundation to transfer the approach of introducing a basic foundation of a taxonomical species concept in primary school to foster further understanding on biodiversity and ecosystem functions.
The timber shortage led to large scale afforestations on previously agriculturally used land in Central Europe during the 19th and 20th century. Widespread afforestation programs created recent forest ecosystems (i.e. young forest systems in terms of their development history). Despite the positive effect of increasing the forest area of Central Europe, the ecological effects of these land-use changes on forest ecosystems remain poorly understood. In order to increase the understanding of ecosystem processes in forests, an assessment of conceivable shifts in ecosystem functions caused by former land-use changes and forest management is required. By analysing aboveground growth rates of European beech (Fagus sylvatica L.) in response to environmental change drivers, such as climate extremes and nitrogen (N) deposition, the presented thesis aims to assess the role of land-use and management legacies in modulating present responses to drivers of environmental change. To this end, annual radial growth rates of individual trees were measured in mature beech stands. The investigated stands differed either in their land-use history (i.e. ancient forest sites versus recent forests) or their forest management history (i.e. managed forest sites versus short-term and long-term unmanaged forest sites). Measurements of radial growth rates were complemented by analyses of the fine root systems, soil chemical properties and crown projection areas to gain insights into the mechanisms underlying alterations in tree growth. Within the projects of the presented thesis, shifts in the climate-growth relationships driven by land-use and management legacies were analysed. In addition, land-use legacy mediated differences in the climate-nitrogen-growth relationships were assessed. The key findings are: (I) Soil legacy driven alterations in the fine root systems cause a higher sensitivity of radial increment rates to water deficits in summer for trees growing on recent forest sites than for trees growing on ancient forest sites. (II) Management legacies (in terms of tree release) enhance the sensitivity of beech’s radial growth to water deficits in spring through changes in crown sizes. (III) Interacting effects of spring water deficits and co-occurring high deposition of reactive N compounds lead to stronger radial growth declines in trees growing in ancient forests. This is likely caused by resource allocation processes towards seed production, which is, in turn, mirrored by decreasing radial growth rates. In this context, high N deposition likely boosts mass fructification in beech trees. Overall, it has been demonstrated that the ecological continuity plays a crucial role in modulating both climate sensitivity and the growth response to interacting effects of water deficits and nitrogen deposition in beech trees. The presented thesis identified a trade-off between the climate sensitivity and maximised growth rates within beech trees, depending on forest history. The results show that the growth of beech in ancient, unmanaged beech forests is less sensitive to water deficits than in recent and managed beech forests. Additionally, interacting effects of spring water deficits and N deposition likely increase the reproductive effort of beech trees, particularly in ancient forests. Thus, the results of this thesis once again underpin the uniqueness of ancient, unmanaged beech forests, whose importance for the conservation of biodiversity has been widely acknowledged. In summary, the presented thesis highlights the need to consider the "ecological memory" of forest ecosystems when predicting responses to current and future environmental changes.
The effects of habitat fragmentation and land use changes are usually studied by relating patterns of genetic diversity and differentiation to environmental factors, habitat history, landscape structure, or to a combination thereof. However, these three drivers are rarely addressed simultaneously. In addition, these studies are usually carried out in conservation-driven contexts, and therefore tend to concentrate on hyper-fragmented landscapes and on rare or endangered species. However, how habitat fragmentation and land use affect widespread species in more typical landscapes has not been fully investigated. This thesis addresses these two gaps. Abax parallelepipedus, a flightless ground beetle with low dispersal power, was used as a model species to test how environmental factors, habitat history, and landscape structure affect genetic diversity and genetic differentiation in three study regions located across Germany. Although all of the study regions represent fairly typical rural landscapes for central Europe, each consisting of a complex matrix of land uses, they differ from one another in terms of environmental factors, habitat history, and landscape structure, and thus can serve as three test cases. In the first stage of the work, the investigator identified polymorphic microsatellite loci which could potentially be used to study genetic diversity and differentiation in A. parallelepipedus. She then developed PCR and genotyping protocols for two suites of loci, in the end selecting to use the set of 14 fully multiplexed loci for the study. After having developed the needed study system, she genotyped over 3300 beetles from 142 study sites. In her investigation of how environmental factors and habitat history affect genetic diversity and genetic differentiation, and found that genetic diversity was being driven by variables that could be related to population sizes rather than by habitat history. She also did not find evidence of an influence of habitat history on the genetic differentiation patterns. Although populations of A. parallelepipedus in the past were probably smaller due to deforestation, they apparently remained large enough to prevent rapid genetic drift. In addition, the researcher carried out a landscape genetics analysis of the genetic differentiation patterns found in each of her study regions, in which she examined the relationship between genetic differentiation and landscape structure. She tested whether she could find patterns of isolation by distance, isolation by resistance, or isolation by barriers in the study regions. No effects of land use or of fragmentation were found. Based on the importance of population sizes found in the previous study, combined with the beetle's known avoidance of non-wooded areas and its inability to cross roads, the investigator concludes that although there is probably little gene flow across the study regions, large population sizes are preventing the rapid development of genetic differentiation. Models simulating the development of genetic differentiation over time in populations of different starting sizes support this conclusion.
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.
Urban areas are prone to climate change impacts. Simultaneously the world's population increasingly resides in cities. In this light, there is a growing need to equip urban decision makers with evidence-based climate information tailored to their specific context to adequately adapt to and prepare for future climate change. To construct climate information high-resolution regional climate models and their projections are pivotal. There is a need to move beyond commonly investigated variables, such as temperature and precipitation, to cover a wider breath of possible climate impacts. In this light, the research presented in this thesis is centered around enhancing the understanding about regional-to-local climate change in Berlin and its surroundings, with a focus on humidity. More specifically, following a regional climate modelling and data analysis approach, this research aims to understand the potential of regional climate models, and the possible added value of convection-permitting simulations, to support the development of high-quality climate information for urban regions, to support knowledge-based decision-making. The first part of the thesis investigates what can already be understood with available regional climate model simulations about future climate change in Berlin and its surroundings, particularly with respect to humidity and related variables. Ten EURO-CORDEX model combinations are analyzed, for the RCP8.5 emission scenario during the time period 1970-2100, for the Berlin region. The results are the first to show an urban-rural humidity contrast under a changing climate, simulated by the EURO-CORDEX ensemble, of around 6% relative humidity, and a robust enlarging urban drying effect, of approximately 2-4% relative humidity, in Berlin compared to its surroundings throughout the 21st century. The second part explores how crossing spatial scales from 12.5km to 3km model grid size affects unprecedented humidity extremes and related variables under future climate conditions for Berlin and its surroundings. Based on the unique HAPPI regional climate model dataset, two unprecedented humidity extremes are identified happening under 1.5°C and 2°C global mean warming, respectively SH>0.02 kg/kg and RH<30%. Employing a double-nesting approach, specifically designed for this study, the two humidity extremes are downscaled to the 12.5km grid resolution with the regional climate model REMO, and thereafter to the 3km with the convection-permitting model version of REMO (REMO NH). The findings indicate that the convection-permitting scale mitigates the SH>0.02kg/kg moist extreme and intensifies the RH<30% dry extreme. The multi-variate process analysis shows that the more profound urban drying effect on the convection-permitting resolution is mainly due to better resolving the physical processes related to the land surface scheme and land-atmosphere interactions on the 3km compared to the 12.5km grid resolution. The results demonstrate the added value of the convection-permitting resolution to simulate future humidity extremes in the urban-rural context. The third part of the research investigates the added value of convection-permitting models to simulate humidity related meteorological conditions driving specific climate change impacts, for the Berlin region. Three novel humidity related impact cases are defined for this research: influenza spread and survival; ragweed pollen dispersion; and in-door mold growth. Simulations by the regional climate model REMO are analyzed for the near future (2041-2050) under emission scenario RCP8.5, on the 12.5km and 3km grid resolution. The findings show that the change signal reverses on the convection-permitting resolution for the impact cases pollen, and mold (positive and negative). For influenza, the convection-permitting resolution intensifies the decrease of influenza days under climate change. Longer periods of consecutive influenza and mold days are projected under near-term climate change. The results show the potential of convection-permitting simulations to generate improved information about climate change impacts in urban regions to support decision makers. Generally, all results show an urban drying effect in Berlin compared to its surroundings for relative and specific humidity under climate change, respectively for the urban-rural contrast throughout the 21st century, for the downscaled future extreme conditions, and for the three humidity related impact cases. Added value for the convection-permitting resolution is found to simulate humidity extremes and the meteorological conditions driving the three impacts cases.
Loss of natural and semi-natural habitat due to increasing human land use for agriculture and housing has led to widespread declines in bee pollinator diversity and abundance, which raised global concerns about the stability of pollination services. Bee population dynamics depend on floral resource diversity and availability in the surrounding landscape, and loss of plant biodiversity may thus directly impair the fitness of individual bee species. However, whether and how plant and resource diversity and availability affect foraging patterns, resource intake, resource quantity and nutrient quality and ultimately fitness of generalist social bees remains unclear. In this thesis, we placed hives of the Australian eusocial stingless bee Tetragonula carbonaria (Apidae, Meliponini) in natural habitat (subtropical forests) and two landscapes differently altered by humans (suburban gardens and macadamia plantations), varying in plant species richness, resource abundance and respective habitat patch size. Foraging patterns and resource intake were compared between landscapes in different seasons and colony growth and fitness were monitored over two and a half years. Bee foraging activity, pollen and sugar intake, diversity of collected pollen and resin resources, resource quantity (colony food stores), colony fitness (brood volume, queenand worker reproduction) and colony growth overwhelmingly increased with plant species richness in the surrounding habitat. However, plant species richness and thus bee fitness was highest in gardens, not in natural forests, as bees in gardens benefited from the continuous floral resource availability of both natural and exotic plants across seasons. In contrast, foraging rates and success, forager orientation and consequently colony fitness was largely reduced in plantations. While bees maximized diversity of collected resources, collecting more diverse resources did however not increase resource functionality and nutritional quality, which appeared to be primarily driven by the surrounding plant community in our study. Conversely, individual worker fitness (body fat and size) was not affected by available resource diversity and abundance, showing that colonies seem not to increase the nutritional investment in single workers, but in overall worker population size. This thesis consequently revealed the outstanding role of plant biodiversity as a key driver of (social) bee fitness by providing more foraging resources, even when only small but florally diverse patches are available.
Ensuring food security and halting biodiversity loss are two of the most pressing global sustainability challenges. Attempts to identify pathways have been dominated with a biophysical-technical focus that provides technical solutions to the integration of food security and biodiversity conservation. The social-political dimension, however, including equity, governance, and empowerment received little to no attention. By focusing on the poorly investigated social-political dimension, this dissertation aimed to identify governance properties that facilitate and impede the integration of food security and biodiversity conservation through an empirical case study conducted in a multi-level governance setting of southwestern Ethiopia. To address the overarching goal of this dissertation, first the author examined how the existing widely discussed food security approaches and agricultural land use framework, land sparing versus land sharing unfold in the local context of southwestern Ethiopia. The finding in this dissertation indicated that the existing global framing of food security approaches as well as frameworks around agricultural land use has limited applicability in on-the-ground realities mainly because landscapes are complex systems that consist of stakeholders with multiple and (often) conflicting interests. This was evident from the finding that local land use preference was not a matter of "either/or", but instead involved mixed features exhibiting properties of both land sparing and land sharing. Moreover, in addition to the biophysical factors embedded in the existing food security approaches and land use frameworks, stakeholders preference involved social factors such as the compatibility of land use strategy with local values and traditions, which are mainly unaccounted in the existing global frameworks. Findings in this dissertation revealed that the existing reductionist analytical framings to the issues of food security and biodiversity conservation seldom address the complexity inherent within and between food security and biodiversity conservation sectors. Second, this dissertation identified governance structural and process related challenges that influence individual as well as integrated achievements of food security and biodiversity conservation goals. The result of the study showed that the governance of food security and biodiversity conservation was characterized by a strongly hierarchical system with mainly linear vertical linkages, lacking horizontal linkages between stakeholders that would transcend administrative boundaries. Furthermore, with regard to the governance process, three key and interdependent categories of governance process challenges namely, institutional misfit, the problem of interplay, and policy incoherence influenced the achievement of individual and integrated goals of food security and were identified. Given the interdependence of these governance challenges, coupled with the complexity inherent in the food security and biodiversity conservation, attempts to achieve the dual goals thus needs an integrative, flexible and adaptive governance system Third, to understand how food security and biodiversity conservation unfold in the future, the author explored future development trajectories for southwestern Ethiopia. Iterative scenario planning process produced four plausible future scenarios that distinctly differed with regard to dominating land use strategies and crops grown, actor constellations and governance mechanisms, and outcomes for food security and biodiversity conservation. Three out of the four scenarios focused on increasing economic gains through intensive and commercial agricultural production. In contrast, one scenario involved features that are widely considered as beneficial to food security and biodiversity conservation, such as agroecological production, diversification practices, and increased social-ecological resilience. In smallholder landscapes such as the one studied here, such a pathway that promises benefits for both food security and biodiversity conservation may need to be given greater emphasis. In order to ensure the integration of food security and biodiversity conservation, recognizing their interdependence and addressing the challenges in a way that fits with the local dynamics is essential. In addition, addressing the food security-biodiversity nexus requires a holistic analytical lens. Moreover, this dissertation indicated that there is a clear need to pay attention to the governance structure that accommodates the diversity of perspectives, enable participation and strong coordination across geographical boundaries, policy domains and governance levels. Finally, this dissertation revealed opportunities to integrate food security and biodiversity through the pro-active management of social-ecological interactions that produce a win-win outcome. The win-win outcome could be achieved in a system that involve properties such as diversification and modern agroecological techniques, smallholders empowerment, emphasize adaptive governance of social-ecological systems, value local knowledge, culture and traditions, and ensure smallholders participation.
Restoration and management of abandoned, dry Continental heathland and sandy grassland communities
(2018)
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. 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. Thus, the present thesis provides substantial knowledge about the population dynamics of the key plant species of dry heaths Calluna vulgaris (L.) HULL 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. 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 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) the author 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. The author 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 the 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. In the third study, the author 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). The author 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 the 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.
Protected areas are an essential tool for conserving biodiversity. However, their ecological effectiveness is contested and their capacity to resist human pressures differ. This dissertation aimed to assess the ecological effectiveness of different protection levels (from strict to less strictly protected: national park, game reserve, forest reserve, game-controlled area, and unprotected areas) in biodiversity (both mega diverse butterflies and mammals), maintaining habitat connectivity, and reducing anthropogenic threats at the wider landscape in the Katavi-Rukwa Ecosystem of southwestern Tanzania. To achieve this overarching goal, the researcher employed an interdisciplinary approach. First, he analyzed butterfly diversity and community composition patterns across protection levels in the Katavi-Rukwa Ecosystem. He found that species richness and abundance were highest in the game reserves and game-controlled areas, intermediate in the forest reserves, national park and unprotected areas. Species composition differed significantly among protection levels. Landscape heterogeneity, forest cover, and primary productivity influenced species composition. Land-use, burned areas, forest cover, and primary productivity explained the richness of species and functional traits. Game reserves hosted most indicator species. Second, the author modelled the spatial distribution of six large mammal target species (buffalo Syncerus caffer, elephant Loxodonta africana, giraffe Giraffa camelopardalis, hartebeest Alcelaphus buselaphus, topi Damaliscus korrigum, and zebra Equus burchellii) across environmental and protection gradients in the Katavi-Rukwa Ecosystem. Based on species-specific density surface models, he found relatively consistent effects of protection level and land-use variables on the spatial distribution of the target mammal species: relative densities were highest in the national park and game reserves, intermediate in forest reserves and game-controlled areas and lowest in un-protected areas. Beyond species-specific environmental predictors for relative densities, the results highlight consistent negative associations between relative densities of the target species and distance to cropland and avoidance of areas in proximity to houses. Third, the author examined temporal changes in land-use, population densities and distribution of six large mammal target species across protection levels between 1991 and 2018. During the surveyed period, cropland increased. Wildlife densities of most, but not all target species declined across the entire landscape. Based on logistic regression models, target species preferred the national park over less strictly protection levels and areas distant to cropland. Fourth, he quantified land-use changes, modelled habitat suitability and connectivity of elephant over time across a large protected area network in southwestern Tanzania. Based on analyses of remotely-sensed data, cropland increased from 7% in 2000 to 13% in 2019. Based on ensemble models, distance from cropland influenced survey-specific habitat suitability for elephant the most. Despite cropland expansion, the locations of the modelled elephant corridors (n=10) remained similar throughout the survey period. Based on circuit theory, the author prioritizes three corridors for protected area connectivity. Key indicators of corridor quality varied over time, whereas elephant movement through some corridors appears to have increased over time. Overall, this dissertation underpins differences in ecological effectiveness of protected areas within one ecosystem. It highlights the need to utilize a landscape conservation approach to guide effective conservation across the entire protection gradient. It also suggests the need to enforcing land use plans and having alternative and sustainable forms for generating income from the land without impairing wildlife habitat.
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”.