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Agricultural land-use may lead to brief pulse exposures of pesticides in edge-of-field streams, potentially resulting in adverse effects on aquatic macrophytes, invertebrates and ecosystem functions. The higher tier risk assessment is mainly based on pond mesocosms which are not designed to mimic stream-typical conditions. Relatively little is known on exposure and effect assessment using stream mesocosms.
Thus the present thesis evaluates the appliacability of the stream mesocosms to mimic stream-typical pulse exposures, to assess resulting effects on flora and fauna and to evaluate aquatic-terrestrial food web coupling. The first objective was to mimic stream-typical pulse exposure scenarios with different durations (≤ 1 to ≥ 24 hours). These exposure scenarios established using a fluorescence tracer were the methodological basis for the effect assessment of an herbicide and an insecticide. In order to evaluate the applicability of stream mesocosms for regulatory purposes, the second objective was to assess effects on two aquatic macrophytes following a 24-h pulse exposure with the herbicide iofensulfuron-sodium (1, 3, 10 and 30 µg/L; n = 3). Growth inhibition of up to 66 and 45% was observed for the total shoot length of Myriophyllum spicatum and Elodea canadensis, respectively. Recovery of this endpoint could be demonstrated within 42 days for both macrophytes. The third objective was to assess effects on structural and functional endpoints following a 6-h pulse exposure of the pyrethroid ether etofenprox (0.05, 0.5 and 5 µg/L; n = 4). The most sensitive structural (abundance of Cloeon simile) and functional (feeding rates of Asellus aquaticus) endpoint revealed significant effects at 0.05 µg/L etofenprox. This concentration was below field-measured etofenprox concentrations and thus suggests that pulse exposures adversely affect invertebrate populations and ecosystem functions in streams. Such pollutions of streams may also result in decreased emergence of aquatic insects and potentially lead to an insect-mediated transfer of pollutants to adjacent food webs. Test systems capable to assess aquatic-terrestrial effects are not yet integrated in mesocosm approaches but might be of interest for substances with bioaccumulation potential. Here, the fourth part provides an aquatic-terrestrial model ecosystem capable to assess cross-ecosystem effects. Information on the riparian food web such as the contribution of aquatic (up to 71%) and terrestrial (up to 29%) insect prey to the diet of the riparian spider Tetragnatha extensa was assessed via stable isotope ratios (δ13C and δ15N). Thus, the present thesis provides the methodological basis to assess aquatic-terrestrial pollutant transfer and effects on the riparian food web.
Overall the results of this thesis indicate, that stream mesocosms can be used to mimic stream-typical pulse exposures of pesticides, to assess resulting effects on macrophytes and invertebrates within prospective environmental risk assessment (ERA) and to evaluate changes in riparian food webs.
Pelagic oxyclines, the transition zone between oxygen rich surface waters and oxygen depleted deep waters, are a common characteristic of eutrophic lakes during summer stratification. They can have tremendous effects on the biodiversity and the ecosystem functioning of lakes and, to add insult to injury, are expected to become more frequent and more pronounced as climate warming progresses. On these grounds, this thesis endeavors to advance the understanding of formation, persistence, and consequences of pelagic oxyclines: We test, whether the formation of metalimnetic oxygen minima is intrinsically tied to a locally enhanced oxygen consuming process, investigate the relative importance of vertical physical oxygen transport and biochemical oxygen consumption for the persistence of pelagic oxyclines, and finally assess their potential consequences for whole lake cycling. To pursue these objectives, the present thesis nearly exclusively resorts to in situ measurements. Field campaigns were conducted at three lakes in Germany featuring different types of oxyclines and resolved either a short (hours to days) or a long (weeks to months) time scale. Measurements comprised temperature, current velocity, and concentrations of oxygen and reduced substances in high temporal and vertical resolution. Additionally, vertical transport was estimated by applying the eddy correlation technique within the pelagic region for the first time. The thesis revealed, that the formation of metalimnetic oxygen minima does not necessarily depend on locally enhanced oxygen depletion, but can solely result from gradients and curvatures of oxygen concentration and depletion and their relative position to each other. Physical oxygen transport was found to be relevant for oxycline persistence when it considerably postponed anoxia on a long time scale. However, its influence on oxygen dynamics was minor on short time scales, although mixing and transport were highly variable. Biochemical consumption always dominated the fate of oxygen in pelagic oxyclines. It was primarily determined by the oxidative breakdown of organic matter originating from the epilimnion, whereas in meromictic lakes, the oxidation of reduced substances dominated. Beyond that, the results of the thesis emphasize that pelagic oxyclines can be a hotspot of mineralization and, hence, short-circuit carbon and nutrient cycling in the upper part of the water column. Overall, the present thesis highlights the importance of considering physical transport as well as biochemical cycling in future studies.
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.
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.
Grundwasser ist eine lebenswichtige Ressource und gleichzeitig ein thermisch stabiler Lebensraum mit einer außergewöhnlichen Fauna. Aufgrund der Klimageschichte der letzten ca. 1,8 Millionen Jahre in Mitteleuropa war die Ausgangshypothese, dass die meisten Grundwasserarten an Temperaturen unterhalb von 14 °C angepasst sind und sich Temperaturerhöhungen daher negativ auf die Grundwasserfauna auswirken.
Um die Folgen einer Klimaerwärmung in situ beurteilen zu können, wurden anthropogen erwärmte Grundwässer als Modelle herangezogen. In ersten Teil dieser Arbeit wurden die Crustaceagemeinschaften natürlich temperierter und anthropogen erwärmter Standorte untersucht. Dazu wurden insgesamt 70 Grundwassermessstellen im Oberrheingraben über ein Jahr (2011 bis 2012) sechsmal beprobt. Ergänzend zu diesem synökologischen Ansatz wurde in einem zweiten Teil die Temperaturpräferenz ausgewählter Arten überprüft. Für diese autökologische Betrachtung wurde ein deutschlandweiter Datensatz herangezogen.
Die ausgewählten Untersuchungsgebiete haben sich als geeignete Modelle für die prognostizierten Temperaturerhöhungen herausgestellt. Die Temperatur erwies sich als einer der wichtigsten Parameter für die Crustaceagemeinschaften. Erwärmungen beeinflussen die Zusammensetzung der Gemeinschaften und die Diversität. Dabei zeigte sich, dass es artspezifisch unterschiedliche Temperaturpräferenzen gibt. Einige vermutlich weniger streng stygobionte Arten scheinen höhere Temperaturen zu tolerieren. Der Großteil der stygobionten Grundwasserarten bevorzugt allerdings Temperaturen unterhalb von 14 °C, wobei wenige Arten als „extrem kalt¬steno-therm“ einzustufen sind. Diese Arten wurden an Standorten nachgewiesen, die aufgrund der hydrologischen Verhältnisse relativ kühl sind und der Fauna auch zukünftig als kalte Refugien Schutz bieten könnten. Einer dieser Standorte ist aufgrund seiner Artenvielalt als „Hot Spot“ einzustufen. Die Ergebnisse legen nahe, dass vor allem kaltstenotherme Arten durch eine Klimaerwärmung und anthropogene Wärmeeinträge gefährdet sind.
Die Untersuchung zeigt, dass Grundwasserlebensräume vor Temperaturerhöhungen zu bewahren sind. Lokale anthropogene Wärmeeinträge sollten keine dauerhafte und großflächige Grundwassererwärmung über 14 °C zur Folge haben. Thermische Ein-träge sollten überwacht und bewertet werden. Es wird empfohlen, die bestehenden rechtlichen Rahmenbedingungen den ökologischen Erfordernissen anzupassen.
Die deutsche Holzhausbranche wächst seit Jahren, allerdings beschäftigen sich die Unternehmen nicht strategisch mit dem Thema Marketing. Diese Dissertation bildet durch qualitative und quantitative Bewohnerbefragungen die Basis für die Entwicklung eines strategischen Marketingansatzes im Holzhausbereich.
Natural pest control and pollination are important ecosystem services for agriculture. They can be supported by organic farming and by seminatural habitats at the local and landscape scale.
The potential of seminatural habitats to support predatory flies (chapters 2 and 3) and bees(chapter 7) at the local and landscape scale was investigated in seminatural habitats. Predatory flies were more abundant in woody habitats and positively related to landscape complexity. The diversity and the abundance of honey and wild bees were positively related to the supply of flowers offered in the seminatural habitats.
The influence of organic farming, adjacent seminatural habitats and landscape complexity on pest control (chapter 4) and pollination (chapter 6) was investigated in 18 pumpkin fields. Organic farming lacked strong effects both on the pest control and on the pollination of pumpkin.
Pest control is best supported at the local scale by the flower abundance in the adjacent habitat. The flower supply positively affected the density of natural enemies and tended to reduce aphid densities in pumpkin fields.
Pumpkin provides a striking example for a dominant role of wild pollinators for pollination success, because bumble bees are the key pollinators of pumpkin in Germany, despite a higher visitation frequency of honey bees. Pollination is best supported by landscape complexity. Bumble bee visits and as a result pollen delivery in pumpkin were negatively related to the dominance of agricultural land in the surrounding landscape.
The influence of aphid density (chapter 8) and pollination (chapter 5) on pumpkin yield was evaluated. Pumpkin yields were not affected by aphid densities observed in the pumpkin fields and not limited by pollination at the current levels of bee visitation.
In conclusion, especially seminatural habitats, that provide diverse, continuous floral resources, are important for natural enemies and pollinators. A sufficient proportion of different seminatural habitat types in agricultural landscapes should be maintained and restored. Thereby natural enemies such as predatory flies, wild pollinators such as bumble bees, and the pest control and pollination provided by them can be supported.
Grassland management has been increasingly intensified throughout centuries since mankind started to control and modify the landscape. Species communities were always shaped alongside management changes leading to huge alterations in species richness and diversity up to the point where land use intensity exceeded the threshold. Since then biodiversity became increasingly lost. Today, global biodiversity and especially grassland biodiversity is pushed beyond its boundaries. Policymakers and conservationists seek for management options which fulfill the requirements of agronomic interests as well as biodiversity conservation alongside with the maintenance of ecosystem processes. However, there is and will always be a trade-off.
Earlier in history, natural circumstances in a landscape mainly determined regionally adapted land use. These regional adaptions shaped islands for many specialist species, and thus diverse species communities, favoring the establishment of a high β-diversity. With the raising food demand, these regional and traditional management regimes became widely unprofitable, and the invention of mineral fertilizers ultimately led to a wide homogenization of grassland management and, as follows, the loss of biotic heterogeneity. In the course of the green revolution, this immediate coherence and the dependency between grassland biodiversity and traditional land use practices becomes increasingly noticed. Indeed, some traditional forms of management such as meadow irrigation have been preserved in a few regions and thus give us the opportunity to directly investigate their long-term relevance for the species communities and ecosystem processes. Traditional meadow irrigation was a common management practice to improve productivity in lowland, but also alpine hay meadows throughout Europe until the 20th century. Nowadays, meadow irrigation is only practiced as a relic in a few remnant areas. In parts of the Queichwiesen meadows flood irrigation goes back to the Middle Ages, which makes them a predestined as a model region to study the long- and short-term effects of lowland meadow irrigation on the biodiversity and ecosystem processes.
Our study pointed out the conservation value of traditional meadow irrigation for the preservation of local species communities as well as the plant diversity at the landscape scale. The structurally more complex irrigated meadows lead to the assumption of a higher arthropod diversity (Orthodoptera, Carabidae, Araneae), which could not be detected. However, irrigated meadows are a significant habitat for moisture dependent arthropod species. In the light of the agronomic potential, flood irrigation could be a way to at least reduce fertilizer costs to a certain degree and possibly prevent overfertilization pulses which are necessarily hazardous to non-target ecosystems. Still, the reestablishment of flood irrigation in formerly irrigated meadows, or even the establishment of new irrigation systems needs ecological and economic evaluation dependent on regional circumstances and specific species communities, at which this study could serve as a reference point.
For a comprehensive understanding of evolutionary processes and for providing reliable prognoses about the future consequences of environmental change, it is essential to reveal the genetic basis underlying adaptive responses. The importance of this goal increases in light of ongoing climate change, which confronts organisms worldwide with new selection pressures and requires rapid evolutionary change to avoid local extinction. Thereby, freshwater ectotherms like daphnids are particularly threatened. Unraveling the genetic basis of local adaptation is complicated by the interplay of forces affecting patterns of genetic divergence among populations. Due to their key position in freshwater communities, cyclic parthenogenetic mode of reproduction and resting propagules (which form biological archives), daphnids are particularly suited for this purpose.
The aim of this thesis was to assess the impact of local thermal selection on the Daphnia longispina complex and to reveal the underlying genetic loci. Therefore, I compared genetic differentiation among populations containing Daphnia galeata, Daphnia longispina and their interspecific hybrids across time, space, and species boundaries. I revealed strongly contrasting patterns of genetic differentiation between selectively neutral and functional candidate gene markers, between the two species, and among samples from different lakes, suggesting (together with a correlation with habitat temperatures) local thermal selection acting on candidate gene TRY5F and indicating adaptive introgression. To reveal the candidate genes’ impact on fitness, I performed association analyses among data on genotypes and phenotypic traits of D. galeata clones from seven populations. The tests revealed a general temperature effect as well as inter-population differences in phenotypic traits and imply a possible contribution of the candidate genes to life-history traits. Finally, utilizing a combined population transcriptomic and reverse ecology approach, I introduced a methodology with a wide range of applications in evolutionary biology and revealed that local thermal selection was probably a minor force in shaping sequence and gene expression divergence among four D. galeata populations, but contributed to sequence divergence among two populations. I identified many transcripts possibly under selection or contributing strongly to population divergence, a large amount thereof putatively under local thermal selection, and showed that genetic and gene expression variation is not depleted specifically in temperature-related candidate genes.
In conclusion, I detected signs of local adaptation in the D. longispina complex across space, time, and species barriers. Populations and species remained genetically divergent, although increased gene flow possibly contributed, together with genotypes recruited from the resting egg bank, to the maintenance of standing genetic variation. Further work is required to accurately determine the influence of introgression and the effects of candidate genes on individual fitness. While I found no evidence suggesting a response to intense local thermal selection, the high resilience and adaptive potential regarding environmental change I observed suggest positive future prospects for the populations of the D. longispina complex. However, overall, due to the continuing environmental degradation, daphnids and other aquatic invertebrates remain vulnerable and threatened.