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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.
Wood-pastures have been present in Europe for thousands of years. This form of grazed landscape, combining herbaceous vegetation with trees and shrubs, has often co-evolved with its human users into complex social-ecological systems (SES). Wood-pastures are associated with high cultural and biodiversity values and are an example of the sustainable use of resources. However, due to their often relatively labour-intensive management and low productivity, large areas of wood-pastures have been lost over the last century. The loss of these areas means not only the loss of biodiversity on both local and landscape scales, but also the loss of traditional farming and cultural heritage in some regions. Across the European Union, wood-pastures are facing different problems and are embedded in different social systems and ecological environments. Yet they are all affected by global change and common European policies. To understand the challenges for wood-pastures in a changing world, a holistic approach combining different disciplines is needed. This dissertation therefore is analyzing wood-pastures across Europe as a Social-ecological System, combining ecology and social science with the aim to identify the barriers and drivers for wood-pastures persistence into the future.
European species-rich grasslands 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 the researchers 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. The results suggest that priority effects can be used by sowing different species or plant functional groups at different time to steer a community to a desired trajectory depending on the restoration goal. However, there is a need to test contingency, potential, and long term impacts of such possible tools for restoration.
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.
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.
In this dissertation, a multi-proxy study, which included palaeoecological, lithological, geochemical and geochronological methods, was carried out to investigate climatic and environmental changes and their interaction during the Quaternary in formerly glaciated and non-glaciated areas. The information obtained will be used to provide a better understanding of the regional stratigraphic framework and to establish broader regional terrestrial correlations within the global marine isotope stage (MIS) framework. This study was conducted on two key drillings, the Garding-2 research drill core in the German North Sea coastal area of Schleswig-Holstein and the GBY#2 archaeological core at the Gesher Benot Ya'aqov (GBY) site, in the Upper Jordan Valley in Israel. The results of this study are presented in three papers. Papers I and II focus on the study of the Garding-2 core, while the multi-proxy study of the GBY#2 core is presented in Paper III. The results of a variety of analyses conducted on the 240 m long Garding-2 sequence show interglacial-glacial cycles that are mainly controlled by variations in temperature. This sequence is composed of mainly fluvial-shallow marine sediments intercalated by muddy-peaty deposits. Based on the palynological and lithological findings, the Pliocene-Pleistocene transition was observed at 182.87 m. It is overlain by Praetiglian and the subsequent sediments of the Waalian and Bavelian Complexes. The boundary of either the second or third Cromerian Interglacial with younger sediments, which still belong to MIS 19, is marked by the last occurrence of Tsuga at 119.50 m and the development of mixed-deciduous forests. The palynologically equivalent sediments of the Bilshausen Interglacial were found below two Elsterian till layers, at 89.00 m-82.00 m. These sediments showed high and increased percentages of Pinus and Picea and scattered occurrences of Abies and Carpinus, which are similar to the features of the beginning of the Bilshausen or Rhume interglacial. An unconformity occurred at 80.29 m, at the bottom of late Holsteinian deposits, characterised by the occurrences of Fagus and Pterocarya, with low percentages of Abies and Carpinus and the absence of Buxus. These deposits are succeeded by sediments of the Fuhne cold period that shows higher percentages of NAP and occurrences of Ericales, Helianthemum and Selaginella selaginoides, which are unconformably overlain by Drenthian till at 73.00 m-71.00 m. A single peaty sample at 69.25 m with Pinus-Picea-Abies assemblage is correlated with the late Eemian Interglacial. This deposit is overlain by Weichselian glaciofluvial sediments. Middle-late Holocene sediments occurred from 20 m upwards, following a hiatus, which was caused by the Early Holocene transgression. A subsequent thin layer of marine Atlantic sediments is unconformably overlain by marine-tidal flat deposits up to 11.00 m. The first occurrence of Fagus (at 15.97 m) and Carpinus (at 15.03 m), which was optically stimulated luminescence (OSL)-dated to 3130 +/- 260 BP (at 16.22 m, Zhang et al., 2014), gives evidence for a Subboreal age for these deposits. Sandy sediments of the early Subatlantic, which were deposited between 11.00 m and the top of the Garding-2 sequence, indicate that local salt marshes, dunes and tidal flat vegetation expanded during this period. Due to regional features and the peculiarities of the local coastal environment, the expansions of Fagus and Carpinus, which are characteristic for the Subboreal-Subatlantic transition at about 2700 BP in northern Germany, are not clearly reflected in the Garding-2 pollen diagram. In the Mediterranean area, a 50 m long core of GBY#2, was drilled at the Acheulian site of Gesher Benot Ya'akov. The GBY#2 core provides a long Early-Middle Pleistocene geological, environmental and climatological record, which also enriches the knowledge of hominin-habitat relationships documented at the margins of the Hula Palaeo-lake. The sediment sequence of GBY#2 is under- and overlain by two basalt flows that are 40Ar/39Ar dated: two samples at the bottom of the core dated to 1195 +/- 67 ka (at 48.30 m) and 1137 +/- 69 ka (at 45.30 m), and another one at the top dated to 659 +/- 85 ka (at 14.90 m). With the additional chronological identification of the Matuyama Brunhes Boundary (MBB) and the correlation with the GBY excavation sites, the sedimentary sequence of GBY#2 provides the climatic history during the late part of the mid-Pleistocene transition (MPT, 1.2 Ma-0.5 Ma). Multi-proxy analyses including those of pollen and non-pollen palynomorphs, macro botanical remains, molluscs, ostracods, fish, amphibians and micromammals provide evidence for lake and lake-margin environments during MIS 20 and MIS 19. During MIS 20, relatively cool semi-moist conditions were followed by a pronounced dry phase. During the subsequent MIS 19, warm and moist interglacial conditions were characterised by Quercus-Pistacia woodlands in this area. The depositional environment changed from an open water lake during MIS 20 to a lake margin environment in MIS 19. This finding is at odds with changing climate conditions from relatively dry to moist. This discrepancy could be explained by the prograding pattern of the lake shore due to the infilling of the basin, which resulted in shallower water. Climatic changes during the Late Tertiary and the Quaternary in the high latitude regions in northwest Europe and during the Early-Middle Pleistocene in the mid latitude regions of the Middle East follow the patterns of global climatic changes, which are mainly controlled by orbital obliquity (+/-41 ka cycle) during the Early Pleistocene and by orbital eccentricity (+/-100 ka cycle) during the MPT (1.2 Ma-0.5 Ma) and the younger periods of the Quaternary. The results of this study also provide reliable evidence for long distance correlation of stratigraphic and climatic events of the Quaternary, which extends knowledge of regional and global impact of climatic fluctuations on the environment.
Social insects like honeybees (Apis mellifera) and stingless bees (Apidae: Meliponini) face a relatively high risk to be attacked by pests and pathogens. To decrease the risk of infection, in addition to an innate immune system, these species have evolved various cooperative defense mechanisms which contribute to the overall health of the colonies and are therefore also referred to as social immunity. The collection and use of plant resin is another important strategy of social immunity. Resin is a sticky, often aromatic substance with antimicrobial and deterrent properties secreted by plants for protection of the vegetative tissue. Honeybees and stingless bees take advantage of these properties by using resins for nest construction (often mixtures of resin and wax called "propolis" or "cerumen") and as defense against pests and pathogens. Plant resins, thus, play a crucial role for the ecology of these species and are an important resource for them. Nevertheless, how bees exploit available resin sources and if resin collection can protect colonies from diseases received comparatively little attention in the past. Therefore the aim of this thesis is to provide new insights into the plant origin and significance as well as the influence of resin resource diversity on bee colony health. For honeybees (A. mellifera) in temperate regions, precise information about which resin sources they use is largely lacking. By chemical comparing bee-collected resins and tree resins, the researcher traced back the resin sources used by individual bees. Results show that honeybees collect distinct resin types that are related to different tree species (several poplar species: Populus balsamifera, P. xcanadensis; Betula alba; Aesculus hippocastanum; several poplar species). With this study, the author provided the first evidence, that A. mellifera in temperate regions use a variety of different tree species as resin sources and, moreover, show preferences for specific resin sources. Furthermore, the researcher conducted an experimental field study in which she investigated whether propolis, as it is naturally deposited in the nests, can protect honeybee colonies against some of the most important pathogens (Varroa destructor mite, Deformed Wing Virus). The results of this study showed that propolis in (semi-) natural conditions can increase the disease resistance of honeybee colonies, underscoring the importance of resins for honeybee health. Resin collection by stingless bees is comparatively well studied and it is known that these species commonly forage on a variety of different plant species. To increase knowledge on whether and how bees may profit from a diversity of resin resources, the researcheer tested how the protective function of a resin varied among different sources (and their mixtures) and various potential aggressors (predators, parasites and pathogens). The results of this study revealed that resins from different trees vary in their effectivity against different target organisms. Moreover, resin blends were more effective than some of the individual resins, suggesting that bees can benefit from a variety of resin resources. In summary, honeybees in temperate regions, similar to tropical stingless bees, use a variety of different tree species as resin sources. Because resins from different tree species varied in their protective function, this indicates that bees can profit from a variety of different resins/resin sources by improving the defense against diverse pests and pathogens. Conversely, the lack of resin had a negative impact on the disease resistance of colonies. Consequently, availability as well as the variety of suitable resin sources is of great importance for the health of bees.
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.
To counteract species loss and preserve the remaining biodiversity, with its important ecosystem functioning and services essential to human well-being, there is an urgent need to develop promising and long-term conservation strategies. In order to achieve these goals, extensive research to gain a better understanding of the general mechanisms underlying community diversity is of greatest importance. Especially, the identification of intrinsic ecological and distributional species traits is receiving increased attention in ecology and conservation biology research. Depending on the expression of their traits, species perform particular ecosystem functions and respond in a specific manner to environmental conditions. Although insects make up the largest part of animal diversity, the majority of studies on extinctions have mainly focused on vertebrates. Among invertebrates either charismatic taxa or those targeted by conservation laws have been investigated until now (e.g. butterflies or saproxylic beetles). Being highly species-rich and trait-diverse, ground beetles (Coleoptera: Carabidae) should be even more suitable for conducting trait-based analyses. Thus, using ground beetles as a model taxon, four case studies focusing on the analyses of traits form the basis of this doctoral thesis. The work of this thesis was conducted with the aim of gaining general insights on the influence of species traits on ground beetle community compositions, such as habitat occupancy and species vulnerability to extinction, for instance. An important aspect when investigating species traits is the consideration of confounding factors, such as dependent relations between the different traits. Compiling a large dataset of 555 Central European species, the author identified that dependent relations between the six tested traits of ground beetles (distribution range size, habitat specialization, body size, hind-wing morphology, breeding season and trophic level) are highly common. Across all identified dependent trait relations, the relation between body size and hind wing morphology or range size and hind wing morphology showed the strongest significant dependencies. Since the consideration of trait relations is necessary to provide reliable interpretations, all analyses of this thesis tested several traits simultaneously and considered possible trait interactions. Studies on local communities found specific traits characterizing the local species pools of certain habitat types. Here, the species pools of seven different habitat types (coastal, forest, mountain, open, riparian, wetland and special habitat) were used to determine habitat-specific trait filters. The identified traits, characteristic for certain habitat types, were in most cases in accordance with the previous findings on local communities. Across Germany, the species of frequently disturbed habitat types, namely coastal, riparian and wetland habitats were characterized by small body size, high amount of macroptery, intermediate to high habitat specialization, spring breeding, and predatory feeding behavior. The species of stable habitat types (forest, mountain, and open habitats), however, were found to be generally larger in body size and more frequently breeding in autumn, further displaying greater variations in the other traits. The gained knowledge on the habitat-specific filtering of traits improve our understanding of the organization and assembly of communities, and can thereby help to detect alterations in the habitat-specific species pool due to natural or human-induced environmental changes. Furthermore, traits can provide evidence on species occurrences and vulnerability to extinction. Three case studies of this thesis aimed to gain new insights on this topic, through the investigations on the following research questions: (1) Which traits drive species extinction risks of Central European ground beetle species? (2) How traits influence the species occurrences of 28 forest species within a large area in Central Europe? (3) Whether certain traits are related to long-term population trends of the species pool from an ancient forest in northern Germany? The results indicated, that depending on the habitat type and tested species pool, different traits prove to be good predictors for the vulnerability of species. Nevertheless, across different geographical and taxonomical scales, especially species with small range sizes and high habitat specialization faced a greater risk of extinction. Therefore, the two traits distributional range size and habitat specialization emerge as reliable predictors of ground beetles vulnerability to extinction. Interestingly, body size did not display a consistent response; while increasing body size led to higher extinction risk in riparian, wetland and open habitats and large macropterous species showed higher extinction risks across the entire species pool, smaller species showed long-term population declines in an ancient forest.
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.