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Vertebrate biodiversity is rapidly decreasing worldwide with amphibians being the most endangered vertebrate group. In the EU, 21 of 89 amphibian species are recognized as being endangered. The intensively used European agricultural landscape is one of the major causes for these declines. As agriculture represents an essential habitat for amphibians, exposure to pesticides can have adverse effects on amphibian populations. Currently, the European risk assessment of pesticides for vertebrates requires specific approaches for fish regarding aquatic vertebrate toxicity and birds as well as mammals for terrestrial vertebrate toxicity but does not address the unique characteristics of amphibians. Therefore, the overall goal of this thesis was to investigate the ecotoxicological effects of pesticides on Central European anuran amphibians. For this, effects on aquatic and terrestrial amphibian life stages as well as on reproduction were investigated. Then, in anticipation of a risk assessment of pesticides for amphibians, this thesis discussed potential regulatory risk assessment approaches.
For the investigated pesticides and amphibian species, it was observed that the acute aquatic toxicity of pesticides can be addressed using the existing aquatic risk assessment approach based on fish toxicity data. However, lethal as well as sublethal effects were observed in terrestrial juveniles after dermal exposure to environmentally realistic pesticide concentrations, which cannot be covered using an existing risk assessment approach. Therefore, pesticides should also be evaluated for potential terrestrial toxicity using risk assessment tools before approval. Additionally, effects of co-formulants and adjuvants of pesticides need specific consideration in a future risk assessment as they can increase toxicity of pesticides to aquatic and terrestrial amphibian stages. The chronic duration of combined aquatic and terrestrial exposure was shown to affect amphibian reproduction. Currently, such effects cannot be captured by the existing risk assessment as data involving field scenarios analysing effects of multiple pesticides on amphibian reproduction are too rare to allow comparison to data of other terrestrial vertebrates such as birds and mammals. In the light of these findings, future research should not only address acute and lethal effects, but also chronic and sublethal effects on a population level. As pesticide exposure can adversely affect amphibian populations, their application should be considered even more carefully to avoid further amphibian declines. Overall, this thesis emphasizes the urgent need for a protective pesticide risk assessment for amphibians to preserve and promote stable amphibian populations in agricultural landscapes.
The role of alternative resources for pollinators and aphid predators in agricultural landscapes
(2021)
The world wide decline of insects is often associated with loss of natural and semi-natural habitat caused by intensified land-use. Many insects provide important ecosystem services to agriculture, such as pest control or pollination. To efficiently promote insects on remaining semi-natural habitat we need precise knowledge of their requirements to non-crop habitat. This thesis focuses on identifying
the most important semi-natural habitats (forest edges, grasslands, and semi-open habitats) for pollinators and natural enemies of crop pests with respect to their food resource requirements. Special
attention is given to floral resources and their spatio-temporal distribution in agricultural landscapes.
Floral resource maps might get closer at characterizing landscapes the way they are experienced by insects compared to classical habitat maps. Performance of the two map types was compared on the prediction of wild bees and natural enemies that consume nectar and pollen, identifying habitats of special importance in the process. In wild bees, influences of spatio-temporal floral resource availability were analysed as well as habitat preferences of specific groups of bees. Understanding dietary needs of natural enemies of crop pests requires additional knowledge on prey use. To this end, ladybird gut contents have been analysed by means of high-throughput sequencing for insight into aphid prey-use.
Results showed, that wild bees were predicted better by floral resource maps compared to classical habitat maps. Forest edge area, as well as floral resources in forest edges had positive effects on abundance and diversity of rare bees and important crop pollinators. Similar patterns were retained for grassland diversity. Especially early floral resources seemed to have positive effects on wild bees. Crops and fruit trees produced a resource pulse in April that exceeded floral resource availability in May and June by tenfold. Most floral resources in forest edges appeared early in the season, with the highest floral density per area. Grasslands provided the lowest amount of floral resources but highest diversity, which was evenly distributed over the season.
Despite natural enemies need for floral resources, classical habitat maps performed better at predicting natural enemies of crop pests compared to floral resource maps. Classical habitat maps revealed a positive effect of forest edge habitat on the abundance of pest enemies, which translated into improved aphid control. Results from gut content analysis reveal high portions of pest aphid species and nettle aphids as well as a broader insight into prey spectra retained from ladybirds collected from sticky traps compared to individuals collected by hand. The aphid specific primer designed for this purpose will be helpful for identifying aphid consumption by ladybirds in future studies.
Findings of this thesis show the potential of floral resource maps for understanding interactions of wild bees and the landscape but also indicate that natural enemies are limited by other resources. I would like to highlight the positive effects of forest edges for different groups of bees as well as natural enemies and their performance on pest control.
Mathematical models of species dispersal and the resilience of metapopulations against habitat loss
(2021)
Habitat loss and fragmentation due to climate and land-use change are among the biggest threats to biodiversity, as the survival of species relies on suitable habitat area and the possibility to disperse between different patches of habitat. To predict and mitigate the effects of habitat loss, a better understanding of species dispersal is needed. Graph theory provides powerful tools to model metapopulations in changing landscapes with the help of habitat networks, where nodes represent habitat patches and links indicate the possible dispersal pathways between patches.
This thesis adapts tools from graph theory and optimisation to study species dispersal on habitat networks as well as the structure of habitat networks and the effects of habitat loss. In chapter 1, I will give an introduction to the thesis and the different topics presented in this thesis. Chapter 2 will then give a brief summary of tools used in the thesis.
In chapter 3, I present our model on possible range shifts for a generic species. Based on a graph-based dispersal model for a generic aquatic invertebrate with a terrestrial life stage, we developed an optimisation model that models dispersal directed to predefined habitat patches and yields a minimum time until these patches are colonised with respect to the given landscape structure and species dispersal capabilities. We created a time-expanded network based on the original habitat network and solved a mixed integer program to obtain the minimum colonisation time. The results provide maximum possible range shifts, and can be used to estimate how fast newly formed habitat patches can be colonised. Although being specific for this simulation model, the general idea of deriving a surrogate can in principle be adapted to other simulation models.
Next, in chapter 4, I present our model to evaluate the robustness of metapopulations. Based on a variety of habitat networks and different generic species characterised by their dispersal traits and habitat demands, we modeled the permanent loss of habitat patches and subsequent metapopulation dynamics. The results show that species with short dispersal ranges and high local-extinction risks are particularly vulnerable to the loss of habitat across all types of networks. On this basis, we then investigated how well different graph-theoretic metrics of habitat networks can serve as indicators of metapopulation robustness against habitat loss. We identified the clustering coefficient of a network as the only good proxy for metapopulation robustness across all types of species, networks, and habitat loss scenarios.
Finally, in chapter 5, I utilise the results obtained in chapter 4 to identify the areas in a network that should be improved in terms of restoration to maximise the metapopulation robustness under limited resources. More specifically, we exploit our findings that a network’s clustering coefficient is a good indicator for metapopulation robustness and develop two heuristics, a Greedy algorithm and a deducted Lazy Greedy algorithm, that aim at maximising the clustering coefficient of a network. Both algorithms can be applied to any network and are not specific to habitat networks only.
In chapter 6, I will summarize the main findings of this thesis, discuss their limitations and give an outlook of future research topics.
Overall this thesis develops frameworks to study the behaviour of habitat networks and introduces mathematical tools to ecology and thus narrows the gap between mathematics and ecology. While all models in this thesis were developed with a focus on aquatic invertebrates, they can easily be adapted to other metapopulations.
Rivers play an important role in the global water cycle, support biodiversity and ecological integrity. However, river flow and thermal regimes are heavily altered in dammed rivers. These impacts are being exacerbated and become more apparent in rivers fragmented by multiple dams. Recent studies mainly focused on evaluating the cumulative impact of cascade reservoirs on flow or thermal regimes, but the role of upstream reservoirs in shaping the hydrology and hydrodynamics of downstream reservoirs remains poorly understood. To improve the understanding of the hydrodynamics in cascade reservoirs, long-term observational data are used in combination with numerical modeling to investigate the changes in flow and thermal regime in three cascade reservoirs at the upper reach of the Yangtze River. The three studied reservoirs are Xiluodu (XLD), Xiangjiaba (XJB) and Three Gorges Reservoir (TGR). In addition, the effects of single reservoir operation (at seasonal/daily time scale) on hydrodynamics are examined in a large tributary of TGR. The results show that the inflow of TGR has been substantially altered by the two upstream reservoirs with a higher discharge in spring and winter and a reduced peak flow in summer. XJB had no obvious contribution to the variations in inflow of TGR. The seasonal water temperature of TGR was also widely affected by the upstream two reservoirs, i.e., an increase in winter and decrease in spring, associated with a delay in water temperature rise and fall. These effects will probably be intensified in the coming years due to the construction of new reservoirs. The study also underlines the importance of reservoir operation in shaping the hydrodynamics of TGR. The seasonal dynamics of density currents in a tributary bay of TGR are closely related to seasonal reservoir operations. In addition, high-frequency water level fluctuations and flow velocity variations were observed in response to periodic tributary bay oscillations, which are driven by the diurnal discharge variations caused by the operation of TGR. As another consequence of operation of cascade reservoirs, the changes in TGR inflow weakened spring thermal stratification and caused warming in spring, autumn and winter. In response to this change, the intrusions from TGR occurred more frequently as overflow and earlier in spring, which caused a sharp reduction in biomass and frequency of phytoplankton blooms in tributary bays of TGR. This study suggests that high-frequency bay oscillations can potentially be used as an efficient management strategy for controlling algal blooms, which can be included in future multi-objective ecological conservation strategies.
Although most plastic pollution originates on land, current research largely remains focused on aquatic ecosystems. Studies pioneering terrestrial microplastic research have adapted analytical methods from aquatic research without acknowledging the complex nature of soil. Meanwhile, novel methods have been developed and further refined. However, methodical inconsistencies still challenge a comprehensive understanding of microplastic occurrence and fate in and on soil. This review aims to disentangle the variety of state-of-the-art sample preparation techniques for heterogeneous solid matrices to identify and discuss best-practice methods for soil-focused microplastic analyses. We show that soil sampling, homogenization, and aggregate dispersion are often neglected or incompletely documented. Microplastic preconcentration is typically performed by separating inorganic soil constituents with high-density salt solutions. Not yet standardized but currently most used separation setups involve overflowing beakers to retrieve supernatant plastics, although closed-design separation funnels probably reduce the risk of contamination. Fenton reagent may be particularly useful to digest soil organic matter if suspected to interfere with subsequent microplastic quantification. A promising new approach is extraction of target polymers with organic solvents. However, insufficiently characterized soils still impede an informed decision on optimal sample preparation. Further research and method development thus requires thorough validation and quality control with well-characterized matrices to enable robust routine analyses for terrestrial microplastics.
This thesis examined two specific cases of point and diffuse pollution, pesticides and salinisation, which are two of the most concerning stressors of Germany’s freshwater bodies. The findings of this thesis were organized into three major components, of which the first component presents the contribution of WWTPs to pesticide toxicity (Chapter 2). The second component focuses on the current and future background salt ion concentrations under climate change with the absence of anthropogenic activities (Chapter 3). Finally, the third major component shows the response of invertebrate communities in terms of species turnover to levels of salinity change, considered as a proxy for human-driven salinisation (Chapter 4).
The European landscape is dominated by intensive agriculture which leads to widespread impact on the environment. The frequent use of agricultural pesticides is one of the major causes of an ongoing decline in flower-visiting insects (FVIs). The conservation of this ecologically diverse assemblage of mobile, flying insect species is required by international and European policy. To counteract the decrease in species numbers and their abundances, FVIs need to be protected from anthropogenic stressors. European pesticide risk assessment was devised to prevent unacceptable adverse consequences of pesticide use on FVIs. However, there is an ongoing discussion by scientists and policy-makers if the current risk assessment actually provides adequate protection for FVI species.
The first main objective of this thesis was to investigate pesticide impact on FVI species. The scientific literature was reviewed to identify groups of FVIs, summarize their ecology, and determine their habitat. This was followed by a synthesis of studies about the exposure of FVIs in their habitat and subsequent effects. In addition, the acute sensitivity of one FVI group, bee species, to pesticides was studied in laboratory experiments.
The second main objective was to evaluate the European risk assessment for possible deficits and propose improvements to the current framework. Regulatory documents were screened to assess the adequacy of the guidance in place in light of the scientific evidence. The suitability of the honey bee Apis mellifera as the currently only regulatory surrogate species for FVIs was discussed in detail.
The available scientific data show that there are far more groups of FVIs than the usually mentioned bees and butterflies. FVIs include many groups of ecologically different species that live in the entire agricultural landscape. Their habitats in crops and adjacent semi-natural areas can be contaminated by pesticides through multiple pathways. Environmentally realistic exposure of these habitats can lead to severe effects on FVI population parameters. The laboratory studies of acute sensitivity in bee species showed that pesticide effects on FVIs can vary greatly between species and pesticides.
The follow-up critical evaluation of the European FVI risk assessment revealed major shortcomings in exposure and effect assessment. The honey bee proved to be a sufficient surrogate for bee species in lower tier risk assessment. Additional test species may be chosen for higher tier risk assessment to account for ecological differences. This thesis shows that the ecology of FVIs should generally be considered to a greater extent to improve the regulatory process. Data-driven computational approaches could be used as alternative methods to incorporate ecological trait data in spatio-temporal scenarios. Many open questions need to be answered by further research to better understand FVI species and promote necessary changes to risk assessment. In general, other FVI groups than bees need to be investigated. Furthermore, comprehensive data on FVI groups and their ecology need to be collected. Contamination of FVI habitat needs to be linked to exposure of FVI individuals and ecologically complex effects on FVI populations should receive increased attention. In the long term, European FVI risk assessment would benefit from shifting its general principles towards more scientifically informed regulatory decisions. This would require a paradigm shift from arbitrary assumptions and unnecessarily complicated schemes to a substantiated holistic framework.
Environmental processes transforming inorganic nanoparticles: implications on aquatic invertebrates
(2020)
Engineered inorganic nanoparticles (EINPs) are produced and utilized on a large scale and will end up in surface waters. Once in surface waters, EINPs are subjected to transformations induced by environmental processes altering the particles’ fate and inherent toxicity. UV irradiation of photoactive EINPs is defined as one effect-inducing pathway, leading to the formation of reactive oxygen species (ROS), increasing EINP toxicity by exerting oxidative stress in aquatic life. Simultaneously, UV irradiation of photoactive EINP alters the toxicity of co-occurring micropollutants (e.g. pesticides) by affecting their degradation. The presence of natural organic matter (NOM) reduces the agglomeration and sedimentation of EINPs, extending the exposure of pelagic species, while delaying the exposure of benthic species living in and on the sediment, which is suggested as final sink for EINPs. However, the joint impact of NOM and UV irradiation on EINP-induced toxicity, but also EINP-induced degradation of micropollutants, and the resulting risk for aquatic biota, is poorly understood. Although potential effects of EINPs on benthic species are increasingly investigated, the importance of exposure pathways (waterborne or dietary) is unclear, along with the reciprocal pathway of EINPs, i.e. the transport back from aquatic to terrestrial ecosystems. Therefore, this thesis investigates: (i) how the presence of NOM affects the UV-induced toxicity of the model EINP titanium dioxide (nTiO2) on the pelagic organism Daphnia magna, (ii) to which extent UV irradiation of nTiO2 in the presence and absence of NOM modifies the toxicity of six selected pesticides in D. magna, (iii) potential exposure pathway dependent effects of nTiO2 and silver (nAg) EINPs on the benthic organism Gammarus fossarum, and (iv) the transport of nTiO2 and gold EINPs (nAu) via the merolimnic aquatic insect Chaetopteryx villosa back to terrestrial ecosystems. nTiO2 toxicity in D. magna increased up to 280-fold in the presence of UV light, and was mitigated by NOM up to 12-fold. Depending on the pesticide, UV irradiation of nTiO2 reduced but also enhanced pesticide toxicity, by (i) more efficient pesticide degradation, and presumably (ii) formation of toxic by-products, respectively. Likewise, NOM reduced and increased pesticide toxicity, induced by (i) protection of D. magna against locally acting ROS, and (ii) mitigation of pesticide degradation, respectively. Gammarus’ energy assimilation was significantly affected by both EINPs, however, with distinct variation in direction and pathway dependence between nTiO2 and nAg. EINP presence delayed C. villosa emergence by up to 30 days, and revealed up to 40% reduced lipid reserves, while the organisms carried substantial amounts of nAu (~1.5 ng/mg), and nTiO2 (up to 2.7 ng/mg). This thesis shows, that moving test conditions of EINPs towards a more field-relevant approach, meaningfully modifies the risk of EINPs for aquatic organisms. Thereby, more efforts need to be made to understand the relative importance of EINP exposure pathways, especially since a transferability between different types of EINPs may not be given. When considering typically applied risk assessment factors, adverse effects on aquatic systems might already be expected at currently predicted environmental EINP concentrations in the low ng-µg/L range.
Gel effect induced by mucilage in the pore space and consequences on soil physical properties
(2020)
Water uptake, respiration and exudation are some of the biological functions fulfilled by plant roots. They drive plant growth and alter the biogeochemical parameters of soil in the vicinity of roots, the rhizosphere. As a result, soil processes such as water fluxes, carbon and nitrogen exchanges or microbial activity are enhanced in the rhizosphere in comparison to the bulk soil. In particularly, the exudation of mucilage as a gel-like substance by plant roots seems to be a strategy for plants to overcome drought stress by increasing soil water content and soil unsaturated hydraulic conductivity at negative water potentials. Although the variations of soil properties due to mucilage are increasingly understood, a comprehensive understanding of the mechanisms in the pore space leading to such variations is lacking.
The aim of this work was to elucidate the gel properties of mucilage in the pore space, i.e. interparticulate mucilage, in order to link changes of the physico-chemical properties in the rhizosphere to mucilage. The fulfilment of this goal was confronted to the three following challenges: The lack of methods for in situ detection of mucilage in soil; The lack of knowledge concerning the properties of interparticulate mucilage; The unknown relationship between the composition and the properties of model substances and root mucilage produced by various species. These challenges are addressed in several chapters.
In a first instance, a literature review picked information from various scientific fields about methods enabling the characterization of gels and gel phases in soil. The variation of soil properties resulting from biohydrogel swelling in soil was named the gel effect. The combined study of water entrapment of gels and gel phases in soil and soil structural properties in terms of mechanical stability or visual structures proved promising to disentangle the gel effect in soil.
The acquired methodical knowledge was used in the next experiments to detect and characterize the properties of interparticulate gel. 1H NMR relaxometry allows the non-invasive measure of water mobility in porous media. A conceptual model based on the equations describing the relaxation of water protons in porous media was developed to integrate the several gel effects into the NMR parameters and quantify the influence of mucilage on proton relaxation. Rheometry was additionally used to assess mucilage viscosity and soil microstructural stability and ESEM images to visualize the network of interparticulate gel. Combination of the results enabled to identify three main interparticulate gel properties: The spider-web effect restricts the elongation of the polymer chains due to the grip of the polymer network to the surface of soil particles. The polymer network effect illustrates the organization of the polymer network in the pore space according to the environment. The microviscosity effect describes the increased viscosity of interparticulate gel in contrast to free gel. The impact of these properties on soil water mobility and microstructural stability were investigated. Consequences on soil hydraulic and soil mechanical properties found in the literature are further discussed.
The influence of the chemical properties of polymers on gel formation mechanism and gel properties was also investigated. For this, model substances with various uronic acid content, degree of esterification and amount of calcium were tested and their amount of high molecular weight substances was measured. The substances investigated included pectic polysaccharides and chia seed mucilage as model polymers and wheat and maize root mucilage. Polygalacturonic acid and low-methoxy pectin proved as non-suitable model polymers for seed and root mucilage as ionic interactions with calcium control their properties. Mucilage properties rather seem to be governed by weak electrostatic interactions between the entangled polymer chains. The amount of high molecular weight material varies considerably depending on mucilage´s origin and seems to be a straight factor for mucilage’s gel effect in soil. Additionally to the chemical characterization of the high molecular weight compounds, determination of their molecular weight and of their conformation in several mucilages types is needed to draw composition-property profiles. The variations measured between the various mucilages also highlight the necessity to study how the specific properties of the various mucilages fulfill the needs of the plant from which they are exuded.
Finally, the integration of molecular interactions in gel and interparticulate gel properties to explain the physical properties of the rhizosphere was discussed. This approach offers numerous perspectives to clarify for example how water content or hydraulic conductivity in the rhizosphere vary according to the properties of the exuded mucilage. The hypothesis that the gel effect is general for all soil-born exudates showing gel properties was considered. As a result, a classification of soil-born gel phases including roots, seeds, bacteria, hyphae and earthworm’s exuded gel-like material according to their common gel physico-chemical properties is recommended for future research. An outcome could be that the physico-chemical properties of such gels are linked with the extent of the gel effect, with their impact on soil properties and with the functions of the gels in soil.
The bio-insecticide Bacillus thuringiensis israelensis (Bti) has worldwide become the most commonly used agentin mosquito control programs that pursue two main objectives: the control of vector-borne diseases and the reduction of nuisance, mainly coming frommosquitoes that emerge in large quantities from seasonal wetlands. The Upper Rhine Valley, a biodiversity hotspot in Germany, has been treated withBti for decades to reduce mosquito-borne nuisance and increase human well-being.Although Btiis presumed to be an environmentally safe agent,adverse effects on wetland ecosystems are still a matter of debate especially when it comes to long-term and indirect effects on non-target organisms. In light of the above, this thesis aims at investigating direct and indirect effects of Bti-based mosquito control on non-target organisms within wetland food chains.Effects were examinedin studies with increasingeco(toxico)logical complexity, ranging from laboratory over mesocosm to field approaches with a focus on the non-biting Chironomidae and amphibian larvae (Rana temporaria, Lissotriton sp.).In addition, public acceptance of environmentally less invasive alternative mosquito control methods was evaluated within surveys among the local population.
Chironomids were the most severely affected non-target aquatic invertebrates. Bti substantially reduced larval and adult chironomid abundances and modified their species composition. Repeated exposures to commonly used Bti formulations induced sublethal alterations of enzymatic biomarkers activityin frog tadpoles. Bti-induced reductions of chironomid prey availability indirectly decreased body size of newts at metamorphosis and increased predation on newt larvae in mesocosm experiments. Indirect effects of severe reductions in midge biomassmight equally be passed through aquatic but also terrestrial food chains influencing predators of higher trophic levels. The majority ofaffectedpeople in the Upper Rhine Valley expressed a high willingness to contributefinancially to environmentally less harmful mosquito control.Alternative approaches could still include Bti applications excepting treatment of ecologically valuable areas. Potentially rising mosquito levels could be counteracted with local acting mosquito traps in domestic and urban areas because mosquito presence was experienced as most annoying in the home environment.
As Bti-based mosquito control can adversely affect wetland ecosystems, its large-scale applications, including nature conservation areas, should be considered more carefully to avoid harmful consequences for the environmentat the Upper Rhine Valley.This thesis emphasizesthe importance to reconsiderthe current practice of mosquito control and encourage research on alternative mosquito control concepts that are endorsed by the local population. In the context ofthe ongoing amphibian and insect declinesfurther human-induced effects onwetlands should be avoided to preserve biodiversity in functioning ecosystems.