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Magmatic continental rifts often constitute the earliest stage of nascent plate boundaries. These extensional tectonic provinces are characterized by ubiquitous normal faulting and volcanic activity; the spatial pattern, the geometry, and the age of these normal faults can help to unravel the spatiotemporal relationships between extensional deformation, magmatism, and long-wavelength crustal deformation of continental rift provinces. This study focuses on the active faulting in the Kenya Rift of the Cenozoic East African Rift System (EARS) with a focus on the mid-Pleistocene to the present-day.
To examine the early stages of continental break-up in the EARS, this thesis presents a time-averaged minimum extension rate for the inner graben of the Northern Kenya Rift (NKR) for the last 0.5 m.y. Using the TanDEM-X digital elevation model, fault-scarp geometries and associated throws are determined across the volcano-tectonic axis of the inner graben of the NKR. By integrating existing geochronology of faulted units with new ⁴⁰Ar/³⁹Ar radioisotopic dates, time-averaged extension rates are calculated. This study reveals that in the inner graben of the NKR, the long-term extension rate based on mid-Pleistocene to recent brittle deformation has minimum values of 1.0 to 1.6 mm yr⁻¹, locally with values up to 2.0 mm yr⁻¹. In light of virtually inactive border faults of the NKR, we show that extension is focused in the region of the active volcano-tectonic axis in the inner graben, thus highlighting the maturing of continental rifting in the NKR.
The phenomenon of focused extension is further investigated with a structural analysis of the youngest volcanic manifestations of the Kenya Rift, their relationship with extensional structures, and their overprint by Holocene faulting. In this context I analyzed the fault characteristics at the ~36 ka old Menengai Caldera and adjacent areas in the Central Kenya Rift using detailed field mapping and a structure-from-motion-based DEM generated from UAV data. In general, the Holocene intra-rift normal faults are dip-slip faults which strike NNE and thus reflect the present-day tectonic stress field; however, inside Menengai caldera persistent magmatic activity and magmatic resurgence overprints these young structures significantly. The caldera is located at the center of an actively extending rift segment and this and the other volcanic edifices of the Kenya Rift may constitute nucleation points of faulting an magmatic extensional processes that ultimately lead into a future stage of magma-assisted rifting.
When viewed at the scale of the entire Kenya Rift the protracted normal faulting in this region compartmentalizes the larger rift depressions, and influences the sedimentology and the hydrology of the intra-rift basins at a scale of less than 100 km. In the present day, most of the fault-bounded sub-basins of the Kenya Rift are hydrologically isolated due to this combination of faulting and magmatic activity that has generated efficient hydrological barriers that maintain these basins as semi-independent geomorphic entities. This isolation, however, was overcome during wetter climatic conditions during the past when the basins were transiently connected. I therefore also investigated the hydrological connectivity of the rift basins during the African Humid Period of the early Holocene, when climate was wetter. With the help of DEM analysis, lake-highstand indicators, radiocarbon dating, and a review of the fossil record, two lake-river-cascades could be identified: one directed southward, and one directed northward. Both cascades connected presently isolated rift basins during the early Holocene via spillovers of lakes and incised river gorges. This hydrological connection fostered the dispersal of aquatic faunas along the rift, and in addition, the water divide between the two river systems represented the only terrestrial dispersal corridor across the Kenya Rift. The reconstruction explains isolated distributions of Nilotic fish species in Kenya Rift lakes and of Guineo-Congolian mammal species in forests east of the Kenya Rift. On longer timescales, repeated episodes of connectivity and isolation must have occurred. To address this problem I participated in research to analyze a sediment drill core from the Koora basin of the Southern Kenya Rift, which provides a paleo-environmental record of the last 1 Ma. Based on this record it can be concluded that at ~400 ka relatively stable environmental conditions were disrupted by tectonic, hydrological, and ecological changes, resulting in increasingly large and frequent fluctuations in water availability, grassland communities, and woody plant cover. The major environmental shifts reflected in the drill core data coincide with phases where volcano-tectonic activity affected the basin. This thesis therefore shows how protracted extensional tectonic processes and the resulting geomorphologic conditions can affect the hydrology, the paleo-environment and the biodiversity of extensional zones in Kenya and elsewhere.
Biodiversity and abundance of wildlife has dramatically declined in agricultural landscapes. Sown, short-lived wildflower (WF) strips along the margins of crop fields are a widespread and often subsidised in agri-environmental schemes, intended to enhance biodiversity, provide refuges for wild plant and arthropod populations and to provide ecosystem services to crops. Meanwhile, WF elements are also criticised, since their functionality decreases with plant succession, the removal of aged WF strip poses an ecological trap for the attracted arthropod populations and only common and mobile species benefit. Further, insects in WF strips are impacted by pesticides from agricultural fields due to shared boundaries with crop fields and by edge effects. The performance of the measure could be improved by combining several WF strips of different successional stages, each harbouring a unique community of plants and arthropods, into persistent, composite WF block, where successional stages exist in parallel. Monitoring data on many taxa in the literature shows, that a third of species are temporarily present in an ageing WF stip, thus offering composite WF blocks should increase cumulative species richness by 28%-39% compared to annual richness in WF strips. Persistence of composite WF blocks would offer reliable refuge for animal and plant populations, also supporting their predators and herbivores. Further, WF blocks have less boundaries to crops compared to WF strips of the same area, and are less impacted by edge effects and pesticides. Policy implications. Here I suggest a change of conservation practice changing from successional WF strips to composite WF blocks. By regular removal and replacement of aged WF strips either within the block (rotational) or at its margins (rolling), the habitat heterogeneity in composite WF block could be perpetuated. Rolling composite WF blocks change locations over years, and the original location can be reconverted to arable land while a nearby WF block is still available to wildlife. A change in agricultural schemes would be necessary, since in some European countries clustered WF strips are explicitly not subsidised.
A large number and wide variety of lake ecosystem models have been developed and published during the past four decades. We identify two challenges for making further progress in this field. One such challenge is to avoid developing more models largely following the concept of others ('reinventing the wheel'). The other challenge is to avoid focusing on only one type of model, while ignoring new and diverse approaches that have become available ('having tunnel vision'). In this paper, we aim at improving the awareness of existing models and knowledge of concurrent approaches in lake ecosystem modelling, without covering all possible model tools and avenues. First, we present a broad variety of modelling approaches. To illustrate these approaches, we give brief descriptions of rather arbitrarily selected sets of specific models. We deal with static models (steady state and regression models), complex dynamic models (CAEDYM, CE-QUAL-W2, Delft 3D-ECO, LakeMab, LakeWeb, MyLake, PCLake, PROTECH, SALMO), structurally dynamic models and minimal dynamic models. We also discuss a group of approaches that could all be classified as individual based: super-individual models (Piscator, Charisma), physiologically structured models, stage-structured models and traitbased models. We briefly mention genetic algorithms, neural networks, Kalman filters and fuzzy logic. Thereafter, we zoom in, as an in-depth example, on the multi-decadal development and application of the lake ecosystem model PCLake and related models (PCLake Metamodel, Lake Shira Model, IPH-TRIM3D-PCLake). In the discussion, we argue that while the historical development of each approach and model is understandable given its 'leading principle', there are many opportunities for combining approaches. We take the point of view that a single 'right' approach does not exist and should not be strived for. Instead, multiple modelling approaches, applied concurrently to a given problem, can help develop an integrative view on the functioning of lake ecosystems. We end with a set of specific recommendations that may be of help in the further development of lake ecosystem models.
A large number and wide variety of lake ecosystem models have been developed and published during the past four decades. We identify two challenges for making further progress in this field. One such challenge is to avoid developing more models largely following the concept of others ('reinventing the wheel'). The other challenge is to avoid focusing on only one type of model, while ignoring new and diverse approaches that have become available ('having tunnel vision'). In this paper, we aim at improving the awareness of existing models and knowledge of concurrent approaches in lake ecosystem modelling, without covering all possible model tools and avenues. First, we present a broad variety of modelling approaches. To illustrate these approaches, we give brief descriptions of rather arbitrarily selected sets of specific models. We deal with static models (steady state and regression models), complex dynamic models (CAEDYM, CE-QUAL-W2, Delft 3D-ECO, LakeMab, LakeWeb, MyLake, PCLake, PROTECH, SALMO), structurally dynamic models and minimal dynamic models. We also discuss a group of approaches that could all be classified as individual based: super-individual models (Piscator, Charisma), physiologically structured models, stage-structured models and traitbased models. We briefly mention genetic algorithms, neural networks, Kalman filters and fuzzy logic. Thereafter, we zoom in, as an in-depth example, on the multi-decadal development and application of the lake ecosystem model PCLake and related models (PCLake Metamodel, Lake Shira Model, IPH-TRIM3D-PCLake). In the discussion, we argue that while the historical development of each approach and model is understandable given its 'leading principle', there are many opportunities for combining approaches. We take the point of view that a single 'right' approach does not exist and should not be strived for. Instead, multiple modelling approaches, applied concurrently to a given problem, can help develop an integrative view on the functioning of lake ecosystems. We end with a set of specific recommendations that may be of help in the further development of lake ecosystem models.
In most biodiversity studies, taxonomic diversity is the measure for the multiplicity of species and is often considered to represent functional diversity. However, trends in taxonomic diversity and functional diversity may differ, for example, when many functionally similar but taxonomically different species co-occur in a community. The differences between these diversity measures are of particular interest in diversity research for understanding diversity patterns and their underlying mechanisms. We analysed a temporally highly resolved 20-year time series of lake phytoplankton to determine whether taxonomic diversity and functional diversity exhibit similar or contrasting seasonal patterns. We also calculated the functional mean of the community in n-dimensional trait space for each sampling day to gain further insights into the seasonal dynamics of the functional properties of the community. We found an overall weak positive relationship between taxonomic diversity and functional diversity with a distinct seasonal pattern. The two diversity measures showed synchronous behaviour from early spring to mid-summer and a more complex and diverging relationship from autumn to late winter. The functional mean of the community exhibited a recurrent annual pattern with the most prominent changes before and after the clear-water phase. From late autumn to winter, the functional mean of the community and functional diversity were relatively constant while taxonomic diversity declined, suggesting competitive exclusion during this period. A further decline in taxonomic diversity concomitant with increasing functional diversity in late winter to early spring is seen as a result of niche diversification together with competitive exclusion. Under these conditions, several different sets of traits are suitable to thrive, but within one set of functional traits only one, or very few, morphotypes can persist. Taxonomic diversity alone is a weak descriptor of trait diversity in phytoplankton. However, the combined analysis of taxonomic diversity and functional diversity, along with the functional mean of the community, allows for deeper insights into temporal patterns of community assembly and niche diversification.
Developments of future scenarios of Antarctic ecosystems are still in their infancy, whilst predictions of the physical environment are recognized as being of global relevance and corresponding models are under continuous development. However, in the context of environmental change simulations of the future of the Antarctic biosphere are increasingly demanded by decision makers and the public, and are of fundamental scientific interest. This paper briefly reviews existing predictive models applied to Antarctic ecosystems before providing a conceptual framework for the further development of spatially and temporally explicit ecosystem models. The concept suggests how to improve approaches to relating species' habitat description to the physical environment, for which a case study on sea urchins is presented. In addition, the concept integrates existing and new ideas to consider dynamic components, particularly information on the natural history of key species, from physiological experiments and biomolecular analyses. Thereby, we identify and critically discuss gaps in knowledge and methodological limitations. These refer to process understanding of biological complexity, the need for high spatial resolution oceanographic data from the entire water column, and the use of data from biomolecular analyses in support of such ecological approaches. Our goal is to motivate the research community to contribute data and knowledge to a holistic, Antarctic-specific, macroecological framework. Such a framework will facilitate the integration of theoretical and empirical work in Antarctica, improving our mechanistic understanding of this globally influential ecoregion, and supporting actions to secure this biodiversity hotspot and its ecosystem services.
Recent declines in biodiversity have given new urgency to questions about the relationship between land-use change, biodiversity and ecosystem processes. Despite the existence of a large body of research on the effects of land use on species richness, it is unclear whether the effects of land use on species richness are principally direct or indirect, mediated by concomitant changes in ecosystem processes. Therefore, we compared the direct effects of land use (fertilization, mowing and grazing) on species richness with indirect ones (mediated via grassland productivity) for grasslands in central Europe. We measured the richness and above-ground biomass in 150 grassland plots in 3 regions of Germany (the so-called Biodiversity Exploratories). We used univariate and structural equation models to examine direct and indirect land-use effects. The direct effects of mowing (-0.37, effect size) and grazing (0.04) intensity on species richness were stronger compared with the indirect effects of mowing (-0.04) and grazing (-0.01). However, the strong negative effect of fertilization (-0.23) on species richness was mainly indirect, mediated by increased productivity compared with the weak direct negative effect (-0.07). Differences between regions in land-use effects showed five times weaker negative effects of mowing (-0.13) in the region with organic soils (Schorfheide-Chorin), strong overall negative effects of grazing (-0.29) for the region with organic soils opposed to a similar strong positive effect (0.30) in the Hainich-Dun region, whereas the Schwabische Alb region displayed a five times weaker positive effect (0.06) only. Further, fertilization effects on species richness were positive (0.03) for the region with organic soils compared to up to 25 times stronger negative effects in the other two regions. Synthesis. Our results clearly show the importance of studying both direct and indirect effects of land-use intensity. They demonstrate the indirect nature, via productivity, of the negative effect of fertilization intensity on plant species richness in the real-world context of management-induced gradients of intensity of fertilization, mowing and grazing. Finally, they highlight that careful consideration of regional environments is necessary before attempting to generalize land-use effects on species diversity.
The concept that diversity promotes reliability of ecosystem function depends on the pattern that community-level biomass shows lower temporal variability than species-level biomasses. However, this pattern is not universal, as it relies on compensatory or independent species dynamics. When in contrast within--trophic level synchronization occurs, variability of community biomass will approach population-level variability. Current knowledge fails to integrate how species richness, functional distance between species, and the relative importance of predation and competition combine to drive synchronization at different trophic levels. Here we clarify these mechanisms. Intense competition promotes compensatory dynamics in prey, but predators may at the same time increasingly synchronize, under increasing species richness and functional similarity. In contrast, predators and prey both show perfect synchronization under strong top-down control, which is promoted by a combination of low functional distance and high net growth potential of predators. Under such conditions, community-level biomass variability peaks, with major negative consequences for reliability of ecosystem function.
Saharan dust input and seasonal upwelling along North-West Africa provide a model system for studying microbial processes related to the export and recycling of nutrients. This study offers the first molecular characterization of prokaryotic particle-attached (PA; > 3.0 mu m) and free-living (FL; 0.2-3.0 mu m) players in this important ecosystem during August 2016. Environmental drivers for alpha-diversity, bacterial community composition, and differences between FL and PA fractions were identified. The ultra-oligotrophic waters off Senegal were dominated by Cyanobacteria while higher relative abundances of Alphaproteobacteria, Bacteroidetes, Verrucomicrobia, and Planctomycetes (known particle-degraders) occurred in the upwelling area. Temperature, proxy for different water masses, was the best predictor for changes in FL communities. PA community variation was best explained by temperature and ammonium. Bray Curtis dissimilarities between FL and PA were generally very high and correlated with temperature and salinity in surface waters. Greatest similarities between FL and PA occurred at the deep chlorophyll maximum, where bacterial substrate availability was likely highest. This indicates that environmental drivers do not only influence changes among FL and PA communities but also differences between them. This could provide an explanation for contradicting results obtained by different studies regarding the dissimilarity/similarity between FL and PA communities and their biogeochemical functions.
Meta-communities of habitat islands may be essential to maintain biodiversity in anthropogenic landscapes allowing rescue effects in local habitat patches. To understand the species-assembly mechanisms and dynamics of such ecosystems, it is important to test how local plant-community diversity and composition is affected by spatial isolation and hence by dispersal limitation and local environmental conditions acting as filters for local species sorting.We used a system of 46 small wetlands (kettle holes)natural small-scale freshwater habitats rarely considered in nature conservation policiesembedded in an intensively managed agricultural matrix in northern Germany. We compared two types of kettle holes with distinct topographies (flat-sloped, ephemeral, frequently plowed kettle holes vs. steep-sloped, more permanent ones) and determined 254 vascular plant species within these ecosystems, as well as plant functional traits and nearest neighbor distances to other kettle holes.Differences in alpha and beta diversity between steep permanent compared with ephemeral flat kettle holes were mainly explained by species sorting and niche processes and mass effect processes in ephemeral flat kettle holes. The plant-community composition as well as the community trait distribution in terms of life span, breeding system, dispersal ability, and longevity of seed banks significantly differed between the two habitat types. Flat ephemeral kettle holes held a higher percentage of non-perennial plants with a more persistent seed bank, less obligate outbreeders and more species with seed dispersal abilities via animal vectors compared with steep-sloped, more permanent kettle holes that had a higher percentage of wind-dispersed species. In the flat kettle holes, plant-species richness was negatively correlated with the degree of isolation, whereas no such pattern was found for the permanent kettle holes.Synthesis: Environment acts as filter shaping plant diversity (alpha and beta) and plant-community trait distribution between steep permanent compared with ephemeral flat kettle holes supporting species sorting and niche mechanisms as expected, but we identified a mass effect in ephemeral kettle holes only. Flat ephemeral kettle holes can be regarded as meta-ecosystems that strongly depend on seed dispersal and recruitment from a seed bank, whereas neighboring permanent kettle holes have a more stable local species diversity.