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Agricultural pesticides, especially insecticides, are an integral part of modern farming. However, these may often leave their target ecosystems and cause adverse effects in non- target, especially freshwater ecosystems, leading to their deterioration. In this thesis, the focus will be on Insect Growth Regulators (IGRs) that can in many ways cause disruption of the endocrine system of invertebrates. Freshwater invertebrates play important ecological, economic and medical roles, and disruption of their endocrine systems may be crucial, considering the important role hormones play in the developmental and reproductive processes in organisms. Although Endocrine Disruption Chemicals (EDCs) can affect moulting, behaviour, morphology, sexual maturity, time to first brood, egg development time, brood size (fecundity), and sex determination in invertebrates, there is currently no agreement upon how to characterize and assess endocrine disruption (ED). Current traditional ecotoxicity tests for Ecological Risk Assessment (ERA) show limitations on generating data at the population level that may be relevant for the assessment of EDCs, which effects may be sublethal, latent and persist for several generations of species (transgenerational).
It is therefore the primary objective of this thesis to use a test method to investigate adverse effects of EDCs on endpoints concerning development and reproduction in freshwater invertebrates. The full life-cycle test over two generations that includes all sensitive life stages of C. riparius (a sexual reproductive organism) allows an assessment of its reproduction and should be suitable for the investigation of long-term toxicity of EDCs in freshwater invertebrates. C. riparius is appropriate for this purpose because of its short life cycle that enables the assessment of functional endpoints of the organism over several generations. Moreover, the chironomid life cycle consists of a complete metamorphosis controlled by a well-known endocrine mechanism and the endocrine system of insects has been most investigated in great detail among invertebrates. Hence, the full life-cycle test with C. riparius provides an approach to assess functional endpoints (e.g. reproduction, sex ratio) that are population-relevant as a useful amendment to the ERA of EDCs. In the laboratory, C. riparius was exposed to environmentally-relevant concentrations of the selected IGRs in either spiked water or spiked sediment scenario over two subsequent generations.
The results reported in this thesis revealed significant effects of the IGRs on the development and the reproduction of C. riparius with the second (F1) generation showing greater sensitivity. These findings indicated for the first time the suitability of multigenerational testing for various groups of EDCs and strongly suggested considering the full life-cycle of C. riparius as an appropriate test method for a better assessment of EDCs in the freshwater environment. In conclusion, this thesis helps to detect additional information that can be extrapolated at population level and, thus, might contribute to better protection of freshwater ecosystems against the risks of Endocrine Disrupting Chemicals (EDCs.) It may furthermore contribute to changes in the ERA process that are necessary for a real implementation of the new European chemical legislation, REACH (Registration, Evaluation Authorization and Restriction of Chemicals). Finally, significant interactions between temperature, chemical exposure and generation were reported for the first time and, may help predict impacts that may occur in the future, in the field, under predicted climate change scenarios.
Statistical eco(-toxico)logy
(2017)
Freshwaters are of immense importance for human well-being.
Nevertheless, they are currently facing unprecedented levels of threat from habitat loss and degradation, overexploitation, invasive species and
pollution.
To prevent risks to aquatic ecosystems, chemical substances, like agricultural pesticides, have to pass environmental risk assessment (ERA) before entering the market.
Concurrently, large-scale environmental monitoring is used for surveillance of biological and chemical conditions in freshwaters.
This thesis examines statistical methods currently used in ERA.
Moreover, it presents a national-scale compilation of chemical monitoring data, an analysis of drivers and dynamics of chemical pollution in streams and, provides a large-scale risk assessment by combination with results from ERA.
Additionally, software tools have been developed to integrate different datasets used in ERA.
The thesis starts with a brief introduction to ERA and environmental monitoring and gives an overview of the objectives of the thesis.
Chapter 2 addresses experimental setups and their statistical analyses using simulations.
The results show that current designs exhibit unacceptably low statistical power, that statistical methods chosen to fit the type of data provide higher power and that statistical practices in ERA need to be revised.
In chapter 3 we compiled all available pesticide monitoring data from Germany.
Hereby, we focused on small streams, similar to those considered in ERA and used threshold concentrations derived during ERA for a large-scale assessment of threats to freshwaters from pesticides.
This compilation resulted in the most comprehensive dataset on pesticide exposure currently available for Germany.
Using state-of-the-art statistical techniques, that explicitly take the limits of quantification into account, we demonstrate that 25% of small streams are at threat from pesticides.
In particular neonicotinoid pesticides are responsible for these threats.
These are associated with agricultural intensity and can be detected even at low levels of agricultural use.
Moreover, our results indicated that current monitoring underestimates pesticide risks, because of a sampling decoupled from precipitation events.
Additionally, we provide a first large-scale study of annual pesticide exposure dynamics.
Chapters 4 and 5 describe software solutions to simplify and accelerate the integration of data from ERA, environmental monitoring and ecotoxicology that is indispensable for the development of landscape-level risk assessment.
Overall, this thesis contributes to the emerging discipline of statistical ecotoxicology and shows that pesticides pose a large-scale threat to small streams.
Environmental monitoring can provide a post-authorisation feedback to ERA.
However, to protect freshwater ecosystems ERA and environmental monitoring need to be further refined and we provide software solutions to utilise existing data for this purpose.
In the new epoch of Anthropocene, global freshwater resources are experiencing extensive degradation from a multitude of stressors. Consequently, freshwater ecosystems are threatened by a considerable loss of biodiversity as well as substantial decrease in adequate and secured freshwater supply for human usage, not only on local scales, but also on regional to global scales. Large scale assessments of human and ecological impacts of freshwater degradation enable an integrated freshwater management as well as complement small scale approaches. Geographic information systems (GIS) and spatial statistics (SS) have shown considerable potential in ecological and ecotoxicological research to quantify stressor impacts on humans and ecological entitles, and disentangle the relationships between drivers and ecological entities on large scales through an integrated spatial-ecological approach. However, integration of GIS and SS with ecological and ecotoxicological models are scarce and hence the large scale spatial picture of the extent and magnitude of freshwater stressors as well as their human and ecological impacts is still opaque. This Ph.D. thesis contributes novel GIS and SS tools as well as adapts and advances available spatial models and integrates them with ecological models to enable large scale human and ecological impacts identification from freshwater degradation. The main aim was to identify and quantify the effects of stressors, i.e climate change and trace metals, on the freshwater assemblage structure and trait composition, and human health, respectively, on large scales, i.e. European and Asian freshwater networks. The thesis starts with an introduction to the conceptual framework and objectives (chapter 1). It proceeds with outlining two novel open-source algorithms for quantification of the magnitude and effects of catchment scale stressors (chapter 2). The algorithms, i.e. jointly called ATRIC, automatically select an accumulation threshold for stream network extraction from digital elevation models (DEM) by assuring the highest concordance between DEM-derived and traditionally mapped stream networks. Moreover, they delineate catchments and upstream riparian corridors for given stream sampling points after snapping them to the DEM-derived stream network. ATRIC showed similar or better performance than the available comparable algorithms, and is capable of processing large scale datasets. It enables an integrated and transboundary management of freshwater resources by quantifying the magnitude of effects of catchment scale stressors. Spatially shifting temporal points (SSTP), outlined in chapter 3, estimates pooled within-time series (PTS) variograms by spatializing temporal data points and shifting them. Data were pooled by ensuring consistency of spatial structure and temporal stationarity within a time series, while pooling sufficient number of data points and increasing data density for a reliable variogram estimation. SSTP estimated PTS variograms showed higher precision than the available method. The method enables regional scale stressors quantification by filling spatial data gaps integrating temporal information in data scarce regions. In chapter 4, responses of the assumed climate-associated traits from six grouping features to 35 bioclimatic indices for five insect orders were compared, their potential for changing distribution pattern under future climate change was evaluated and the most influential climatic aspects were identified (chapter 4). Traits of temperature preference grouping feature and the insect order Ephemeroptera exhibited the strongest response to climate as well as the highest potential for changing distribution pattern, while seasonal radiation and moisture were the most influential climatic aspects that may drive a change in insect distribution pattern. The results contribute to the trait based freshwater monitoring and change prediction. In chapter 5, the concentrations of 10 trace metals in the drinking water sources were predicted and were compared with guideline values. In more than 53% of the total area of Pakistan, inhabited by more than 74 million people, the drinking water was predicted to be at risk from multiple trace metal contamination. The results inform freshwater management by identifying potential hot spots. The last chapter (6) synthesizes the results and provides a comprehensive discussion on the four studies and on their relevance for freshwater resources conservation and management.
Today’s agriculture heavily relies on pesticides to manage diverse pests and maximise crop yields. Despite elaborate regulation of pesticide use based on a complex environmental risk assessment (ERA) scheme, the widespread use of these biologically active compounds has been shown to be a threat to the environment. For surface waters, pesticide exposure has been observed to exceed safe concentration levels and negatively impact stream ecology leading to the question whether current ERA schemes ensure a sustainable use of pesticides. To answer this, the large-scale “Kleingewässer-Monitoring” (KgM) assessed the occurrence of pesticides and related effects in 124 streams throughout Germany, Central Europe, in 2018 and 2019.
Based on five scientific publications originating from the KgM, this thesis evaluated pesticide exposure in streams, ecological effects and the regulatory implications. More than 1,000 water samples were analysed for over 100 pesticide analytes to characterise occurrence patterns (publication 1). Measured concentrations and effects were used to validate the exposure and effect concentrations predicted in the ERA (publication 2). By jointly analysing real-world pesticide application data and measured pesticide mixtures in streams, the disregard of environmental pesticide mixtures in the ERA was evaluated (publication 3). The toxic potential of mixtures in stream water was additionally investigated using suspect screening for 395 chemicals and a battery of in-vitro bioassays (publication 4). Finally, the results from the KgM stream monitoring were used to assess the capability to identify pesticide risks in governmental monitoring programmes (publication 5).
The results of this thesis reveal the widespread occurrence of pesticides in non-target stream ecosystems. The water samples contained a variety of pesticides occurring in complex mixtures predominantly in short-term peaks after rainfall events (publications 1 & 4). Respective pesticide concentration maxima were linked to declines in vulnerable invertebrate species and exceeded regulatory acceptable concentrations in about 80% of agricultural streams, while these thresholds were still estimated partly insufficient to protect the invertebrate community (publication 2). The co-occurrence of pesticides in streams led to a risk underestimated in the single substance-oriented ERA by a factor of about 3.2 in realistic worst-case scenarios, which is further exacerbated by a high frequency at which non-target organism are exposed to pesticides (publication 3). Stream water samples taken after rainfall caused distinct effects in bioassays which were only explainable to a minor extent by the many analytes, indicating the relevance of unknown chemical or biological mixture components (publication 4). Finally, the regulatory monitoring of surface waters under the Water Framework Directive (WFD) was found to significantly underestimate pesticide risks, as about three quarters of critical pesticides and more than half of streams at risk were overlooked (publication 5).
Essentially, this thesis involves a new level of validation of the ERA of pesticides in aquatic ecosystems by assessing pesticide occurrence and environmental impacts at a scale so far unique. The overall results demonstrate that the current agricultural use of pesticides leads to significant impacts on stream ecology that go beyond the level tolerated under the ERA. This thesis identified the underestimation of pesticide exposure, the potential insufficiency of regulatory thresholds and the general inertia of the authorisation process as the main causes why the ERA fails to meet its objectives. To achieve a sustainable use of pesticides, the thesis proposes substantial refinements of the ERA. Adequate monitoring programmes such as the KgM, which go beyond current government monitoring efforts, will continue to be needed to keep pesticide regulators constantly informed of the validity of their prospective ERA, which will always be subject to uncertainty.
Streams are coupled with their riparian area. Emerging insects from streams can be an important prey in the riparian area. Such aquatic subsidies can cause predators to switch prey or increase predator abundances. This can impact the whole terrestrial food web. Stressors associated with agricultural land use can alter insect communities in water and on land, resulting in complex response patterns of terrestrial predators that rely on prey from both systems.
This thesis comprises studies on the impact of aquatic nsects on a terrestrial model ecosystem (Objective 1, hapter 2), the influence of agricultural land use on riparian spiders’ traits and community (Objective 2, Chapter 3), and on the impact of agricultural land use on the contribution of different prey to spider diet (Objective 3, Chapter 4).
In chapter 2, I present a study where we conducted a mesocosm experiment to examine the effects of aquatic subsidies on a simplified terrestrial food web consisting of two types of herbivores (leafhoppers and weevils), plants and predators (spiders). I focused on the prey choice of the spiders by excluding predator immigration and reproduction. In accordance with predator switching, survival of leafhoppers increased in the presence of aquatic subsidies. By contrast, the presence of aquatic subsidies indirectly reduced weevils and herbivory.
In chapter 3, I present the results on the taxonomic and trait response of riparian spider communities to gradients of agricultural stressors and environmental variables, with a particular emphasis on pesticides. To capture spiders with different traits and survival strategies, we used multiple collection methods. Spider community composition was best explained by in-stream pesticide toxicity and shading of the stream bank, a proxy for the quality of the habitat. Species richness and the number of spider individuals, as well as community ballooning ability, were negatively associated with in-stream pesticide toxicity. In contrast, mean body size and shading preference of spider communities responded strongest to shading,
whereas mean niche width (habitat preference for moisture and shading) responded strongest to other environmental variables.
In chapter 4, I describe aquatic-terrestrial predator-prey relations with gradients of agricultural stressors and environmental variables. I sampled spiders, as well as their aquatic and terrestrial prey along streams with an assumed pesticide pollution gradient and determined their stable carbon and nitrogen signals. Potential aquatic prey biomass correlated positively with an increasing aquatic prey contribution of T. montana. The contribution of aquatic prey to the diet of P. amentata showed a positive relationship with increasing toxicity in streams.
Overall, this thesis contributes to the emerging discipline of cross-ecosystem ecology and shows that aquatic-terrestrial linkages and riparian food webs can be influenced by land use related stressors. Future manipulative field studies on aquatic-terrestrial linkages are required that consider the quality of prey organisms, fostering mechanistic understanding of such crossecosystem effects. Knowledge on these linkages is important to improve understanding of consequences of anthropogenic stressors and to prevent further losses of ecosystems and their biodiversity.
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.
World’s ecosystems are under great pressure satisfying anthropogenic demands, with freshwaters being of central importance. The Millennium Ecosystem Assessment has identified anthropogenic land use and associated stressors as main drivers in jeopardizing stream ecosystem functions and the
biodiversity supported by freshwaters. Adverse effects on the biodiversity of freshwater organisms, such as macroinvertebrates, may propagate to fundamental ecosystem functions, such as organic matter breakdown (OMB) with potentially severe consequences for ecosystem services. In order to adequately protect and preserve freshwater ecosystems, investigations regarding potential and observed as well as direct and indirect effects of anthropogenic land use and associated stressors (e.g. nutrients, pesticides or heavy metals) on ecosystem functioning and stream biodiversity are needed. While greater species diversity most likely benefits ecosystem functions, the direction and magnitude of changes in ecosystem functioning depends primarily on species functional traits. In this context, the functional diversity of stream organisms has been suggested to be a more suitable predictor of changes in ecosystem functions than taxonomic diversity.
The thesis aims at investigating effects of anthropogenic land use on (i) three ecosystem functions by anthropogenic toxicants to identify effect thresholds (chapter 2), (ii) the organic matter breakdown by three land use categories to identify effects on the functional level (chapter 3) and (iii)on the stream community along an established land-use gradient to identify effects on the community level.
In chapter 2, I reviewed the literature regarding pesticide and heavy metal effects on OMB, primary production and community respiration. From each reviewed study that met inclusion criteria, the toxicant concentration resulting in a reduction of at least 20% in an ecosystem function was standardized based on laboratory toxicity data. Effect thresholds were based on the relationship between ecosystem functions and standardized concentration-effect relationships. The analysis revealed that more than one third of pesticide observations indicated reductions in ecosystem functions at concentrations that are assumed being protective in regulation. However, high variation within and between studies hampered the derivation of a concentration-effect relationship and thus effect thresholds.
In chapter 3, I conducted a field study to determine the microbial and invertebrate-mediated OMB by deploying fine and coarse mesh leaf bags in streams with forested, agricultural, vinicultural
and urban riparian land use. Additionally, physicochemical, geographical and habitat parameters were monitored to explain potential differences in OMB among land use types and sites. Regarding results, only microbial OMB differed between land use types. The microbial OMB showed a negative relationship with pH while the invertebrate-mediated OMB was positively related to tree cover. OMB responded to stressor gradients rather than directly to land use.
In chapter 4, macroinvertebrates were sampled in concert with leaf bag deployment and after species identification (i) the taxonomic diversity in terms of Simpson diversity and total taxonomic
richness (TTR) and (ii) the functional diversity in terms of bio-ecological traits and Rao’s quadratic entropy was determined for each community. Additionally, a land-use gradient was established and the response of the taxonomic and functional diversity of invertebrate communities along this gradient was investigated to examine whether these two metrics of biodiversity are predictive for the rate of OMB. Neither bio-ecological traits nor the functional diversity showed a significant relationship with
OMB. Although, TTR decreased with increasing anthropogenic stress and also the community structure and 26 % of bio-ecological traits were significantly related to the stress gradient, any of these shifts propagated to OMB.
Our results show that the complexity of real-world situations in freshwater ecosystems impedes the effect assessment of chemicals and land use for functional endpoints, and consequently our potential to predict changes. We conclude that current safety factors used in chemical risk assessment may not be sufficient for pesticides to protect functional endpoints. Furthermore, simplifying real-world stressor gradients into few land use categories was unsuitable to predict and quantify losses in OMB. Thus, the monitoring of specific stressors may be more relevant than crude land use categories to detect effects on ecosystem functions. This may, however, limit the large scale assessment of the status of OMB. Finally, despite several functional changes in the communities the functional diversity over several trait modalities remained similar. Neither taxonomic nor functional diversity were suitable predictors of OMB. Thus, when understanding anthropogenic impacts on the linkage between biodiversity and ecosystem functioning is of main interest, focusing on diversity metrics that are clearly linked to the stressor in question (Jackson et al. 2016) or integrating taxonomic and functional metrics (Mondy et al., 2012) might enhance our predictive capacity.
Change of ecosystems and the associated loss of biodiversity is among the most important environmental issues. Climate change, pollution, and impoundments are considered as major drivers of biodiversity loss. Organism traits are an appealing tool for the assessment of these three stressors, due to their ability to provide mechanistic links between organism responses and stressors, and consistency over wide geographical areas.
Additionally, traits such as feeding habits influence organismal performance and ecosystem processes. Although the response of traits of specific taxonomic groups to stressors is known, little is known about the response of traits of different taxonomic groups to stressors. Additionally, little is known about the effects of small impoundments on stream ecosystem processes, such as leaf litter decomposition, and food webs.
After briefly introducing the theoretical background and objectives of the studies, this thesis begins by synthesizing the responses of traits of different taxonomic groups to climate change and pollution. Based on 558 peer-reviewed studies, the uniformity (i.e., convergence) in trait response across taxonomic groups was evaluated through meta-analysis (Chapter 2). Convergence was primarily limited to traits related to tolerance.
In Chapter 3, the hypothesis that small impoundments would modify leaf litter decomposition rates at the sites located within the vicinity of impoundments, by altering habitat variables and invertebrate functional feeding groups (FFGs) (i.e., shredders), was tested. Leaf litter decomposition rates were significantly reduced at the study sites located immediately upstream (IU) of impoundments, and were significantly related to the abundance of invertebrate shredders.
In Chapter 4, the invertebrate FFGs were used to evaluate the effect of small impoundments on stream ecosystem attributes. The results showed that heterotrophic production was significantly reduced at the sites IU. With regard to food webs, the contribution of methane gas derived carbon to the biomass of chironomid larvae was evaluated through correlation of stable carbon isotope values of chironomid larvae and methane gas concentrations.
The results indicated that the contribution of methane gas derived carbon into stream benthic food web is low. In conclusion, traits are a useful tool in detecting ecological responses to stressors across taxonomic groups, and the effects of small impoundments on stream ecological integrity and food web are limited.