570 Biowissenschaften; Biologie
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We are in a phase of an alarming biodiversity loss, by scientist already referred to as Earth’s sixth mass extinction. According to estimations, the current extinction rates are 100 to 1000 times higher than those predicted from fossil records. 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. The identification of the effect of certain traits on community compositions can therefore significantly improve our understanding of species extinction processes and help to develop valuable and appropriate recommendations for conservation management. As trait-based analyses are applicable to different geographical, temporal and taxonomical scales, they may even allow for a broader generalization if similar results are found on different scales, i.e. for local species pools, the complete species pools of different habitat types or the entire species pool across several habitat types including different climatic regions. Although insects make up the largest part of animal diversity and provide essential ecosystem services in form of e.g. pollination, pest control, and decomposition, 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 which could influence the results, such as dependent relations between the different traits. Compiling a large dataset of 566 Central European species, I 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; I. Which traits drive species extinction risks of Central European ground beetle species, II. How traits influence the species occurrences of 28 forest species within a large area in Central Europe, and III. 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. To summarize, this thesis presents a comprehensive picture of ground beetle species traits, providing valuable insights and a better understanding of the mechanisms driving changes in ground beetle diversity. On the basis of the results presented in this work, the efficiency of biodiversity protection can be increased by developing appropriate management and recovery plans, especially targeting species of threatened habitat types or ‘functional groups’ of species, exhibiting trait values strongly associated with a greater vulnerability to extinction.
Forest ecosystems significantly contribute to global carbon (C) sequestration and therefore play a crucial role for climate change mitigation. At the same time, forests were and are subjected to past and current environmental changes with consequences for the functioning of forest ecosystems and their associated ecosystem services. Forests in Central Europe are highly influenced by former settlement activities and land-use changes, as well as silvicultural management measures. Until the beginning of the 19th century anthropogenic activities caused a tremendous decline of the forest area. The resulting timber shortage led to large scale afforestations on previously agriculturally used land (e.g. heathlands, grasslands and croplands) 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 addition, most forests in Central Europe are under silvicultural management, while the knowledge about the consequences of management measures on forest ecosystem functioning, particularly in the face of ongoing global environmental changes, is also still limited. 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 with a forest continuity > 230 years versus recent forests afforested on former arable land ~ 100 years ago) 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.
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 such as hygienic behavior or allo-grooming, 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. Resource use and availability form fundamental prerequisites, having decisive influence on the viability of individuals and maintenance of populations. Information on the resources required by a species is thus important to effectively promote and preserve it. 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, I 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 I 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. Maintenance of colony health is probably one of the major purposes of resin collection. Nevertheless, studies investigating the benefits of resins at the colony level are rare and there are only few evidences on the effects of raw propolis (unlike commonly used ethanol extracts) on colony health. For this reason, I conducted an experimental field study in which I 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, I 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. In addition to nectar and pollen, resin, as a further important resource, should therefore find more attention in beekeeping. Resin collection as the natural disease defense of bees should find more respect in beekeeping praxis and should be more strongly included in future consideration on how to promote bee colony health.
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.
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.
The Ili Delta in Kazakhstan is an important ecosystem that offers crucial wetland habitats for several bird species. However, the Ili River, the Ili Delta and the Balkhash Lake are suffering from water shortage due to climate change and human activities. The desertification of the Aral Sea, an obvious point of comparison to the Balkhash region, also involved the degradation of wetland habitats and the related loss of many bird species relying on these habitats. Therefore, water shortage at the Ili Delta may also be the reason for the loss of wetland habitats and bird species. In this study, bird species numbers, species abundances as well as bird diversity at different habitats in the Ili Delta were examined. There are many habitat types provided by the Ili Delta, for example reed bed vegetation, Tugay forest, bare soil floodplains along rivers and steppe. The results of this study showed that the central delta region with habitats of submerged reed vegetation showed the highest number of bird species and the greatest diversity. Threatened bird species at the Ili Delta were also observed only in these wetland habitats. Steppe habitats showed the lowest numbers of bird species and the lowest bird diversity. In general, all habitats at the Ili Delta are important for the ecosystem and essential for the bird species that depend on them for their survival. With expansion of arid steppe habitats due to water shortage, however, previous wetland habitats may be lost. Moreover, bird species that depend on these wetland habitats may also be lost. Therefore, protective measures for the Balkhash region in general and the wetland habitats at the Ili Delta and its distinct avifauna in particular are urgently needed.
The role of tree diversity for individual tree growth, crown architecture and branch demography
(2012)
In the light of the concurrent loss of biodiversity, biodiversity and ecosystem functioning (BEF) research attracted a great deal of attention and emerged as one of the important fields of research in ecology. Since important ecological interactions such as competition occur between individuals, the understanding of individual tree growth was considered to be fundamental for forest related BEF research. Individual tree growth is determined by the above- and belowground interactions of a tree individual with its local neighbourhood. To obtain a deeper understanding of BEF relationships, I broadened the focus from individual tree growth (usually measured as diameter or biomass increment) to the arrangement and dynamics of the above-ground modules of trees in dependence of their local neighbourhood. More precisely, the main objective of the present thesis was to analyse the impact of tree diversity on individual tree growth, crown architectural and branch demographic variables. Thereby I considered crown architectural variables as important indicators of the competition for light. In addition, crown architectural variables impacted ecosystem services such as erosion control. Furthermore, the results of the present thesis contributed to the current discussion on species coexistence theories, which may be differentiated by two opposing views: one that relies on neutral processes and one that implicates a role for meaningful differences in the ecological strategy (niche) of co-occurring species. The studied forest ecosystems were the subtropical broad-leaved evergreen forests of southeast China, which have been under high human pressure due to a long history of intensive land-use. The area is of particular interest for BEF research due to the high species richness of woody plants, including many, yet poorly studied species, and due to the rough terrain with steep slopes, which cause severe soil erosion. The present thesis combines three observational with two experimental studies, applying the local neighbourhood approach along an age gradient from tree saplings to mature trees. In the Gutianshan National Nature Reserve (GNNR), I conducted two observational studies on permanent plots which were chosen according to a space-for-time substitution design. The aim of the first study was to reveal the effects of diversity (species richness, functional diversity) together with other biotic and abiotic variables on morphological growth parameters (crown area, crown displacement and stem inclination) of target trees of four tree species (Castanea henryi, Castanopsis eyrei, Quercus serrata and Schima superba). In the second study, the same target trees together with their neighbours were used to analyse the relation between stand related functional diversity and the horizontal and vertical structure of the canopy. The third study was conducted in a young secondary broad-leaved evergreen forest. Using two target species (Castanopsis fargesii and Quercus fabri), the role of diversity, intra- vs. inter-specific competition and the mode of competition (symmetric vs. asymmetric) on the target individuals was tested by analysing five-year radial growth increments. The two other studies were carried out in an experimentally established plantation, using saplings of four tree species (C. henryi, Elaeocarpus decipiens, Q. serrata and S. superba), which were planted in monoculture, twoand four-species combinations and in three densities. The fourth study focused on mechanisms of coexistence and the role of species richness, species composition, species identity and density on sapling growth. The fifth study tested the effect of sapling density and identity on the througfall kinetic energy, which represents a measure for the erosive power of rain. It was found that functional diversity does affect crown architectural and canopy related parameters of forests in the GNNR. However, no effects of species richness on radial-growth were detected in the younger forest. Since I also did not find strong effects of species richness on saplings in the experimental plantation, diversity effects may evolve at a later age stage. The importance of the diversity effect may be related reversely to that of species identity in an age gradient of forest stands. The findings suggest that different mechanisms of coexistence operate simultaneously but that their relative importance may shift through the life stages of trees. During the sapling stage, species-specific differences in growth and architectural traits support niche theory. In older forest stands, no species-specific differences in growth parameters could be detected. However, I did find effects of functional diversity on horizontal canopy structure. I conclude that mechanisms of coexistence may not only change with forest stand age, but may also differ for distinct traits. The present thesis, being the first to apply the local neighbourhood approach with regard to crown architecture and branch demography within the BEF field of research, stresses the importance of this individual based approach. Although the observed forest systems are very complex, crown architectural and canopy structural variables were found to be affected by diversity. The finding that the degree of erosive power of rain could be elucidated by crown architectural variables, encourages further studies to reveal possible relations between biodiversity and other ecosystem functions or services, which might be mediated by crown architectural and canopy structural variables.