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Institute
Leaf litter breakdown is a fundamental process in aquatic ecosystems, being mainly mediated by decomposer-detritivore systems that are composed of microbial decomposers and leaf-shredding, detritivorous invertebrates. The ecological integrity of these systems can, however, be disturbed, amongst others, by chemical stressors. Fungicides might pose a particular risk as they can have negative effects on the involved microbial decomposers but may also affect shredders via both waterborne toxicity and their diet; the latter by toxic effects due to dietary exposure as a result of fungicides’ accumulation on leaf material and by negatively affecting fungal leaf decomposers, on which shredders’ nutrition heavily relies. The primary aim of this thesis was therefore to provide an in-depth assessment of the ecotoxicological implications of fungicides in a model decomposer-detritivore system using a tiered experimental approach to investigate (1) waterborne toxicity in a model shredder, i.e., Gammarus fossarum, (2) structural and functional implications in leaf-associated microbial communities, and (3) the relative importance of waterborne and diet-related effects for the model shredder.
Additionally, knowledge gaps were tackled that were related to potential differences in the ecotoxicological impact of inorganic (also authorized for organic farming in large parts of the world) and organic fungicides, the mixture toxicity of these substances, the field-relevance of their effects, and the appropriateness of current environmental risk assessment (ERA).
In the course of this thesis, major differences in the effects of inorganic and organic fungicides on the model decomposer-detritivore system were uncovered; e.g., the palatability of leaves for G. fossarum was increased by inorganic fungicides but deteriorated by organic substances. Furthermore, non-additive action of fungicides was observed, rendering mixture effects of these substances hardly predictable. While the relative importance of the waterborne and diet-related effect pathway for the model shredder seems to depend on the fungicide group and the exposure concentration, it was demonstrated that neither path must be ignored due to additive action. Finally, it was shown that effects can be expected at field-relevant fungicide levels and that current ERA may provide insufficient protection for decomposer-detritivore systems. To safeguard aquatic ecosystem functioning, this thesis thus recommends including leaf-associated microbial communities and long-term feeding studies using detritus feeders in ERA testing schemes, and identifies several knowledge gaps whose filling seems mandatory to develop further reasonable refinements for fungicide ERA.
Population genetic structure in European Hyalodaphnia species: Monopolization versus gene flow
(2012)
Cyclic parthenogens displays an alternation of asexual and sexual reproduction which has consequences for the genetic structure of these organisms. The clonal diversity of cyclic parthenogenetic zooplankton populations is influenced by the size of the dormant egg bank, i.e., the amount of sexually produced dormant eggs that assembled in the sediment, as these dormant eggs contribute new genetic variants to the populations. Further, the clonal diversity is impacted by clonal erosion over time, which reduces the number of different clones through stochastic and selective processes. Although freshwater invertebrates are good dispersers through their dormant stages, the influence of gene flow is assumed to be negligible, as the local population successfully monopolizes the available resources. As these populations reach carrying capacity fast due to the asexual reproduction, the first colonizing individuals are able to successfully establish in the habitat, resulting in a priority effect which hinders the invasion of new genotypes. Due to clonal selection and sexual reproduction a population will locally adapt over time and will establish a dormant egg bank which facilitates the fast re-colonization after a hostile period. This thesis evaluates the processes altering the population genetic structure of cyclic parthenogenetic zooplankton with a special focus on the concepts of monopolization as well as the counteracting effects of gene flow, using large-lake Daphnia species. Thirty-two variable microsatellite DNA markers were developed and a subset of twelve markers was evaluated regarding their suitability for species assignment and hybrid class detection. With this marker set and an additional mitochondrial DNA marker forty-four natural European populations of the species D. cucullata, D. galeata and D. longispina were studied. In D. galeata, most populations were characterized by low clonal diversities which suggest high influence from clonal erosion over the growing season and a low contribution from the dormant egg bank. Further, recent expansions as well as gene flow were detected, probably caused by the anthropogenic alteration of freshwater habitats, in particular eutrophication of many European lakes. D. longispina and D. cucullata revealed a different genetic structure compared to D. galeata, with high genetic differentiation among populations. This indicates low levels of effective gene flow which is in line with the predictions of monopolization. Further, high clonal diversities were found in populations of the two taxa, suggesting a high contribution from the dormant egg bank while clonal erosion was often not detectable. In D. longispina, mitochondrial data revealed an ancient expansion which was probably initiated by the formation of glacial lakes after the last ice age.
In addition, in D. longispina not only clonal diversity but also genetic diversity was high, indicating that during the build-up of the studied populations the influence from gene flow was probably high. To better understand the processes that act on early populations the population build-up in regard to the temporal advantage of clones during invasion succession was experimentally studied and revealed that priority effects shape population structure of Daphnia species. However, in certain cases the highly superior clones resulted in the extinction of inferior clones independent of the temporal advantage the single clones had.
This clearly shows that not only the time of succession is important but also the competitive strength. rnIn conclusion, the results obtained show that the population genetic structure in cyclic parthenogenetic zooplankton species is impacted by various processes. In addition to earlier studies, which mainly focus on local adaptation, clonal erosion and the size of the dormant egg bank to understand population genetic structure, this thesis could show that gene flow may be effective as well. During population build-up the advantage of early arriving individuals does not necessarily predict the outcome of population assembly, as additional genotypes may contribute to the population. Finally, the genetic structure of established populations may be severely impacted by effective gene flow, if severe environmental changes alter the habitat of the locally adapted population.
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.
The conservation of groundwater ecosystems requires an assessment and evaluation scheme that shows the state of the ecosystem. Consequently, faunal and microbial criteria are required for groundwater monitoring, in addition to physico-chemical analyses. To proof the adequacy of the application of groundwater organisms for the biological assessment of groundwater quality, an extensive sampling collection of various groundwater systems was accomplished between 2002 and 2009 in Germany. Key aspects were the examination of the indicator potential of groundwater communities towards surface-groundwater interactions and anthropogenic impacts, as well as the analysis of stygofaunal distribution patterns, as a base and reference for a faunal assessment scheme of groundwater systems. The sampling design considered local, regional, and biogeographic conditions. To test the indicator potential of groundwater organisms on a local and regional scale, groundwater systems in Nordrhein-Westfalen (NRW) and Baden-Württemberg (BW) were examined. The faunal and microbial data from unconsolidated aquifers (Erftgebiet, NRW) show the sensitivity of groundwater organisms against land use effects, e.g. extensive agriculture. Data analyses revealed positive correlations of organic material (estimated amount of detritus, TOC) and nitrate with faunal abundance, species richness, diversity and the proportions of stygoxene species. Moreover, the bacterial abundance and diversity in the groundwater of the Erftgebiet was high compared to oligotrophic groundwater systems, indicating an effect of surface influence due to agricultural land use. The groundwater colonization in the Alb-Donau-Kreis (BW) was analyzed for regional effects (landscape, type of aquifer, hydrogeology) and local effects (comparing single wells). The results show that the fauna reflects the strength of the hydrological exchange on different spatial scales. Furthermore, the fauna reflects the interaction of regional and local conditions. Accordingly, the diversity and abundance of groundwater organisms was influenced by the high connectivity of the karst and unconsolidated alluvial aquifers, the type of land use, covering layers of soil, the age of groundwater, and the sampling depths. In general, faunal and microbial data of the Alb-Donau-Kreis are characteristic for oligotrophic, oligoalimonic groundwaters. The large scale analyses of stygofaunal distribution patterns revealed significant biogeographical differences of the communities. These community patterns of the groundwater fauna do not coincide with existing classification schemes defined for surface landscapes or freshwater systems. The largest differences between faunal surface and subsurface distribution patterns were found between the groundwater of northern and southern Germany and the foothills of the central mountain ranges - all of them regions shaped by the last ice ages.
In accordance with the faunal data assessed in groundwater, four different stygoregions were defined that are populated by distinct faunal assemblages. These are 1) the "Northern lowlands", 2) the "Central mountain ranges", 3) the "South-western mountain ranges", and 4) the "Southern mountain ranges and northern alps". The study corroborates that stygofaunal and microbial communities are an adequate tool for the qualitative assessment and monitoring of groundwater ecosystems. The best indicators to detect anthropogenic impacts on groundwater ecosystems are the faunal diversity, abundance, the proportions of stygobitic (obligate groundwater species) and stygoxene species (species not obligate for groundwater), and the bacterial abundance. The development of an ecologically based groundwater assessment and management, is crucial for the conservation of our groundwater ecosystems and thus, healthy drinking water. The defintion of "stygoregions" is an important base for the development of an assessment and reference scheme for groundwater ecosystems. The assessment of the ecological state of groundwater systems must be conducted on local scale, because the strength of the local hydrological influence determines the amount of food and oxygen as well as stygoxene animals within the aquifer, and thus shapes the local groundwater communities. Nevertheless, information of the biogeographic and regional conditions is needed as a reference for the type of species and abundances that potentially can occur in certain groundwater systems.
Assessment of bat activity in agricultural environments and the evaluation of the risk of pesticides
(2013)
Although agriculture dominates with around 50% area much of Europe- landscape, there is virtually no information on how bats use this farmed environment for foraging. Consequently, little is known about effective conservation measures to compensate potential negative effects of agrarian management practice on the food availability for bats in this habitat. Moreover, there are currently no specific regulatory requirements to include bats in European Union risk assessments for the registration of pesticides since no information about pesticide exposure on this mammal group is available. To evaluate the potential pesticide exposure of bats via ingestion of contaminated insects, information about bat presence and activity in agricultural habitats is required. In order to examine bat activity on a landscape scale it was necessary to establish a suitable survey method. Contrary to capture methods, telemetry, and direct observations, acoustic surveys of bat activity are a logistically feasible and cost-effective way of obtaining bat activity data. However, concerns regarding the methodological designs of many acoustic surveys are expressed in the scientific literature. The reasons are the failing of addressing temporal and spatial variation in bat activity patterns and the limitations of the suitability of the used acoustic detector systems. By comparing different methods and detector systems it was found that the set up of several stationary calibrated detector systems which automatically trigger the ultrasonic recording has the highest potential to produce reliable, unbiased and comparable data sets on the relative activity of bats.
By using the proposed survey method, bat diversity and activity was recorded in different crops and semi-natural habitats in southern Rhineland-Palatinate. Simultaneously, the availability of aerial prey insects was studied by using light and sticky traps. In more than 500 sampling nights about 110,000 call sequences were acoustically recorded and almost 120,000 nocturnal insects were sampled. A total of 14 bat species were recorded, among them the locally rare and critically endangered northern bat (Eptesicus nilssonii) and the barbastelle (Barbastella barbastellum), all of them also occurring over agricultural fields. The agricultural landscape of southern Palatinate is dominated by vineyards, a habitat that was shown to be of low quality for most bat species because of the demonstrated low availability of small aerial insects. By surveying bat activity and food availably in a pair-wise design on several rain water retention ponds and neighbouring vineyards it was demonstrated that aquatic insect emergence in artificial wetlands can provide an important resource subsidy for bats. The creation of artificial wetlands would be a possibility to create important foraging habitats for bats and mitigate negative effects of management practice in the agricultural landscape.
In several other agricultural crops, however, high abundances of suitable prey insects and high bat activity levels, comparable or even higher than in the nearby forests and meadows known to be used as foraging habitats were demonstrated. Especially high bat activity levels were recorded over several fruit orchards and vegetable fields where insects were also present. Both crops are known for high pesticide inputs, and, therefore, a pesticide exposure through ingestion of contaminated insects can not be excluded. To follow the current risk assessment approach for birds and mammals pesticide residues were measured on bat-specific food items in an apple orchard following insecticide applications and bat activity was recorded in parallel. The highest residue values were measured on foliage-dwelling arthropods which may results in a reproductive risk for all bat species that, even to a small extent, include this prey group in their diet. The presence of bats in agricultural landscapes that form a majority of the land area in Europe but also on a global scale leads to exposure of bats by contaminated food and depletion of their food resources by pesticide use. So far conservation efforts for bats focussed on securing hibernation sites and the creation of artificial roost sites since especially the latter were thought to be limiting population growth. However the potential pesticide effects might be also crucial for the population persistence in agricultural landscapes of bats and need to be addressed adequately, especially in risk assessment procedures for the regulation of pesticides.
This habilitation thesis deals with the effects of toxicants on freshwater ecosystems and considers different toxicant classes (pesticides, organic toxicants, salinity) and biotic endpoints (taxonomic community structure, trait community structure, ecosystem functions).
The thesis comprises 12 peer-reviewed international publications on these topics. All of the related studies rely on mesocosm or field investigations, or the analysis of field biomonitoring or chemical monitoring data. Publications I and II are devoted to passive sampling of a neonicotinoid insecticide and polycyclic aromatic hydrocarbons (PAHs), respectively. They show that biofouling and a diffusion-limiting membrane can reduce the sampling rate of the pulsed insecticide exposure and that receiving phases of different thicknesses can be used to assess the kinetic regime during field deployment of passive samplers. Publications III to VI mainly focus on trait-based approaches to reveal toxicant effects on invertebrates in streams. An overview on the framework and several applications of a trait-based approach to detect effects of pesticides (SPEARpesticides index) are given in publication III. Publication IV describes the development of a trait database for South-East Australian stream invertebrates and its successful application in the adaptation of SPEARpesticides as well as the development of a salinity index. Moreover, a conceptual model for the future development of trait-based biomonitoring indices is proposed. Publication V reports a mesocom study on the effects of a neonicotinoid insecticide on field-realistic invertebrate communities. The insecticide had long-term effects on the invertebrate communities, which were only detected when grouping the taxa according to their life-history traits. A comprehensive field study employing different pesticide sampling methods including passive sampling and biomonitoring of the invertebrate and microbial communities is presented in publication VI. The study did not find pesticide-induced changes in the microbial communities, but detected adverse effects of current-use pesticides on the invertebrate communities using the trait-based SPEARpesticides index. This index is also applied in a meta-analysis on thresholds for the effects of pesticides on invertebrate communities in publication VII. It is shown that there is a similar dose-response relationship between SPEARpesticides and pesticide toxicity over different biogeographical regions and continents. In addition, the thresholds for effects of pesticides are lower than derived from most mesocosm studies and than considered in regulatory pesticide risk assessment. The publications VIII to X use statistical data analysis approaches to examine effects of toxicants in freshwater ecosystems. Using governmental monitoring data on 331 organic toxicants monitored monthly in 4 rivers over 11 years, publication VIII finds that organic toxicants frequently occurred in concentrations envisaging acute toxic effects on invertebrates and algae even in large rivers. Insecticides and herbicides were the chemical groups mainly contributing to the ecotoxicological risk. Publication IX introduces a novel statistical method based on a similarity index to estimate thresholds for the effects of toxicants or other stressors on ecological communities. The application of the method for deriving thresholds for salinity, heavy metals and pesticides in streams is presented in three case studies. Publication X tackles the question of interactive effects between different toxicants using data from a field study on stream invertebrates in 24 sites of South-East Australia. Both salinity and pesticides exhibited statistically significant effects on the invertebrate communities, but no interaction between the stressors was found. Moreover, salinity acted on a higher taxonomical level than pesticides suggesting evolutionary adaptation of stream invertebrates compared to pesticide stress. Publications XI and XII concentrate on the effects of toxicants on biodiversity, ecosystem functions and ecosystem services, with publication XI summarising different studies related to the ecological risk assessment for these endpoints. A field study on the effects of pesticides and salinity on the ecosystem functions of allochthonous organic matter decomposition, gross primary production and ecosystem respiration is presented in publication XII. Both pesticides and salinity reduced the breakdown of allochthonous organic matter, whereas no effects on the other ecosystem functions were detected. A chapter following these publications synoptically discusses all studies of this habilitation thesis and draws general conclusions. It is stressed that in order to advance the understanding of effects of toxicants on freshwater ecosystems more ecological realism is needed in ecotoxicological approaches and that the spatiotemporal extent of toxicant effects needs more scrutiny.
Worldwide one third to one half of the freshwater crayfish species are threatened with population decline or extinction. Besides habitat deterioration, pollution, and other man-made environmental changes, invasive species and pathogens are major threats to the survival of European crayfish species. Freshwater crayfish are the largest freshwater invertebrates and strongly influence the structure of food webs. The disappearance of crayfish from a water body may change the food web and could have dramatic consequences for an ecosystem.rnOne goal in modern species conservation strategies is the conservation of genetic diversity, since genetic diversity is an advantage for the long-term survival of a species. The main aim of my thesis was to reveal the genetic structure and to identify genetic hotspots of the endangered noble crayfish (Astacus astacus) throughout Europe (part 1 of my thesis). Since the most significant threat to biodiversity of European crayfish species is the crayfish plague pathogen Aphanomyces astaci I studied new aspects in the distribution of A. astaci (part two of my thesis). The results serve as a basis for future conservation programs for freshwater crayfish. In the first part of my thesis I conducted a phylogeographic analysis of noble crayfish using mitochondrial DNA and nuclear microsatellite data. With these methods I aimed to identify its genetic hotspots and to reconstruct the recolonization history of central Europe by this species. I detected high genetic diversities in southestern Europe indicating that noble crayfish outlasted the cold climate phases during the Pleistocene in this region (Appendix 1). Because of the high genetic diversity found there, southeastern Europe is of particular importance for the conservation of noble crayfish. The mitochondrial DNA analysis points to a bifurcated colonization process from the eastern Black Sea basin to a) the North Sea and to b) the Baltic Sea basin (Appendix 2). A second independent refugium that was localized on the Western Balkans did not contribute to the colonization of central Europe. Furthermore, I found that the natural genetic structure is dissolved, probably due to the high human impact on the distribution of noble crayfish (e.g. artificial translocation). In the second part of this thesis using real-time PCR I identified calico crayfish (Orconectes immunis) as the fourth North American crayfish species to be carrier of the agent of the crayfish plague (Appendix 3). Furthermore I detected the crayfish plague pathogen in American spiny-cheek crayfish (Orconectes limosus) and native narrow-clawed crayfish (Astacus leptodactylus) in the lower Danube in Romania (Appendix 4). The distribution of infected spiny-cheek crayfish poses a threat to the native biodiversity in southeastern Europe and shows the high invasion potential of this crayfish species. Moreover, I found that even the native narrow-clawed crayfish in the Danube Delta, about 970 km downstream of the current invasion front of American crayfish, is a carrier of A. astaci (Appendix 5). This finding is of high importance, as the native species do not seem to suffer from the infection. In Appendix 6 I elucidate demonstrate that the absence of the crayfish plague agent is the most likely explanation for the coexistence of populations of European and American crayfish in central Europe. In my thesis I show that the common assumption that all North American crayfish are carrier of A. astaci and that all native crayfish species die when infected with A. astaci does not hold true. The studies presented in my thesis reveal new aspects that are crucial for native crayfish conservation: 1) The genetic diversity of noble crayfish is highest in southeastern Europe where noble crayfish outlasted the last glacial maximum in at least two different refugia. 2) Not all American crayfish populations are carrier of A. astaci and 3) not all Europen crayish populations die shortly after being infected with the crayfish plague pathogen.rnTo conserve native crayfish species and their (genetic) diversity in the long term, further introductions of American crayfish into European waters must be avoided. However, the introduction will only decrease if the commercial trade with non-indigenous crayfish species is prohibited.
To assess the effect of organic compounds on the aquatic environment, organisms are typically exposed to toxicant solutions and the adverse effects observed are linked to the concentration in the surrounding media. As compounds generally need to be taken up into the organism and distributed to the respective target sites for the induction of effects, the internal exposure is postulated to best represent the observed effects.
The aim of this work is to contribute to an improved effect assessment of organic compounds by describing experimental and modelling methods to obtain information on the internal exposure of contaminants in organisms.
Chapter 2 details a protocol for the determination of bioconcentration parameter for uptake (k1) and elimination (k2) of organic compounds in zebrafish (Danio rerio) eggs. This enables the simulation of the internal exposure in zebrafish eggs from an ambient exposure concentration over time. The accumulated contaminant amount in zebrafish eggs was also determined, using a biomimetic extraction method. Different bioconc-entration estimation models for the determination of internal steady-state concentrat-ion of pharmaceutical compounds in fish to an environmental exposure are presented in Chapter 3. Bioconcentration factors were estimated from the compounds octanol: water partition coefficient (KOW) to determine the internal exposure to an ambient concentration.
To assess the integral bioavailable fraction from the water and sediment phase of environmental contaminants for rooted aquatic plants, the internal exposure in river-living Myriophyllum aquaticum plants were determined over time, presented in Chapter 4. The plants were collected at different time points, with the accumulated organic contaminants determined using a liquid extraction method.
In Chapter 5 a protocol was established to enable the non-invasive observation of effects in M. aquaticum plants exposed to contaminated sediments over time. Since the toxicant effects are a result of all uptake and distribution processes to the target site and the toxico-dynamic process leading to an observed effect during static exposure, information on the internal exposure could thus be gained from the temporal effect expression.rn
Field margins are often the only remaining habitats of various wild plant species in agricultural landscapes. However, due to their proximity to agricultural fields, the vegetation of field margins can be affected by agrochemicals applied to the crop fields. The aim of this thesis was to investigate the individual and combined effects of herbicide, insecticide and fertilizer inputs on the plant community of a field margin. Therefore, a 3-year field experiment with a randomized block design including seven treatments (H: herbicide, I: insecticide, F: fertilizer, H+I, F+I, F+H and F+H+I) and one control was conducted on a low-production meadow. Each treatment was replicated 8 times in 8 m x 8 m plots with a distance of 2 m between each plot. The fertilizer rates (25 % of the field rate) and pesticide rates (30 % of the field rate) used for the plot applications were consistent with realistic average input rates (overspray + drift) in the first meter of a field margin directly adjacent to a wheat field.
The study revealed that fertilizer and herbicide misplacements in field margins are major factors that affect the natural plant communities of these habitats. In total, 20 of the 26 abundant species on the study site were significantly affected by the fertilizer and herbicide treatment. The fertilizer promoted plants with high nutrient uptake and decreased the frequencies of small species. The herbicide caused a nearly complete disappearance of three species directly after the first application, whereas sublethal effects (e.g., phytotoxic effects and reduced seed productions of up to 100 %) were observed for the other affected species. However, if field margins are exposed to repeated agrochemical applications over several years, then such sublethal effects (particularly reproduction effects) also reduce the population size of plant species significantly, as observed in this study.
Significant herbicide-fertilizer interaction effects were also detected and could not be extrapolated from individual effects. The fertilizer and herbicide effects became stronger over time, leading to shifts in plant community compositions after three years and to a 15 % lower species diversity than in the control. The insecticide significantly affected the frequencies of two plant species (1 positively and 1 negatively). The results of the experiment suggest that a continuous annual agrochemical application on the study site would cause further plant community shifts and would likely lead to the disappearance of certain affected plants. A clear trend of increasing grass dominance at the expense of flowering herbs was detected. This finding corresponds well with monitoring data from field margins near the study site.
Although herbicide risk assessment aims to protect non-target plants in off-field habitats from adverse effects, reproduction effects and combined effects are currently not considered. Furthermore, no regulations for fertilizer applications next to field margins exist and thus, fertilizer misplacements in field margins are likely to occur and to interact with herbicide effects.
Adaptations of the current risk assessment, a development of risk mitigation measures (e.g., in-field buffers) for the application of herbicides and fertilizers, and general management measures for field margins are needed to restore and conserve plant diversity in field margins in agricultural landscapes.
Quellen sind hochdiverse Lebensräume mit einer stark spezialisierten Fauna, wobei noch Lücken im Verständnis der besiedlungsrelevanten Faktoren für die aquatische Makrofauna bestehen, so dass die Arbeit einen klärenden Beitrag hinsichtlich anthropogener Veränderungen und ihrer Auswirkungen auf die Besied-lung leisten will. Basierend auf Daten von 334 Quellen aller Grundwasserlandschaften von Rheinland-Pfalz wurden die Quellen morphologisch und faunistisch bewertet. Außerdem wurden die Quellen des Landes morphologisch typisiert, Referenzquellen analysiert und ein Ansatz für die morphologische und faunistische Leitbildentwicklung gegeben. Außerdem wird ein selbst entwickeltes, kompaktes Kartier- und Bewertungsverfahren zur Quellstruktur vorgestellt, wo bislang noch ein Defizit bestand. Das Verfahren erfasst in einem Erfassungsbogen alle ökologisch bedeutsamen Strukturparameter und bewertet in einem 5-stufigen System. Quellen sind hochdiverse Lebensräume mit kleinen Populationen in isolierter Lage. Bei der Untersuchung der 310 Quellen wurden insgesamt 292 Arten bzw. höhere Taxa des Makrozoobenthos nachgewiesen. Es wurden 89 quellassoziierte Taxa gezählt, hiervon waren 24 krenobiont und 65 krenophil. Der Anteil der quellassoziierten Fauna an der Gesamtfauna lag bei über 30%. Es fanden sich 19 Erstnachweise für Rheinland-Pfalz, 9 besondere Nachweise (seltene Arten) und 19 Rote-Liste-Arten. Erstnachweise fanden sich in wenig bearbeiteten Dipterengruppen, vor allem den Psychodidae (13), Limoniidae (5) und ferner den Simu-liidae (1). Bei den Köcherfliegen wurden etliche seltene Arten gefunden, welche bisher in Rheinland-Pfalz kaum gefunden wurden. Häufigere krenobionte Taxa waren Crunoecia irrorata, Bythinella dunkeri, Niphar-gus sp., Pisidium personatum, Salamandra salamandra, Thaumalea sp., Agabus guttatus, Crenobia alpina, Oxycera sp. und Beraea maura. Die durchschnittliche Taxazahl pro Quelle betrug 11, die maximale 62 Taxa. Schüttung und Morphologie der Quellen waren von Naturraum und Grundwasserlandschaft abhängig, so schütteten etwa Buntsandsteinquellen ergiebiger und konstanter als Tonschieferquellen. Quellen spiegeln die Hydrologie ihrer Einzugsgebiete wider, so dass in verschiedenen hydrogeologischen Einheiten die chemischen Parameter differierten. In einzelnen Quellen schwankte die Hydrochemie jahreszeitlich aber nur gering. Der pH-Wert etwa war im Quellwasser versauerungsgefährdeter Gebiete niedrig (Hunsrück, Pfälzer-wald), der Nitratgehalt anthropogen bedingt hoch in intensiv landwirtschaftlich genutzten Räumen. Die morphologische Quelltypologie, welche sich in erster Linie auf gängige Quelltypen stützte, ergab in elf hydrogeologischen Quelltypenräumen vier Basisquelltypen und fünf geochemische Sondertypen. Die Wan-derquelle wurde neu hinzu genommen. Die landesweite Verteilung der Quelltypen ergab bei ungefassten Quellen 57 % Sicker-, 32 % Sturz-, 9 % Tümpel- und über 2 % Wanderquellen. Wanderquellen dürften aber noch öfter vorkommen. Relief und Substrateigenschaften waren entscheidende Faktoren für die Typologie, so dass etwa Sturzquellen vor allem im Buntsandstein häufig waren, während Sickerquellen im Tonschiefer des rheinischen Schiefergebirges dominierten. Es wurden 85 Strukturreferenzquellen als typische, naturnahe Beispiele für einzelne Quelltypenräume genannt. Etwa zwei Drittel aller Quellen waren strukturell beein-trächtigt bis stark geschädigt, 59 % gefasst. Fassungen, Verrohrungen und Betonverbau waren ökologisch besonders bedeutsam. In Regionen mit intensiver Landwirtschaft lagen generell die stärksten Schädigungen vor. Im Forst waren Quellen durch Nadelholzmonokulturen und in den Hochlagen pufferarmer Mittelgebirge durch Versauerung geschädigt. Hinsichtlich der Besiedlung ergaben sich Unterschiede zwischen den Naturräumen, beeinflusst durch die Faktoren Geologie, Hydrochemie, Schüttungsverhältnisse, Landnutzung im Einzugsgebiet sowie die Struk-turvielfalt an der Quelle. Strukturell unveränderte und unversauerte Quellen besaßen im Schnitt höhere Ta-xazahlen. Einzelne Faktoren mit Einfluss auf die Besiedlung wurden in ihrer Auswirkung getestet. In Quel-len ist die Versauerung grundsätzlich besiedlungsbeschränkend und überlagert strukturelle Faktoren. Bei niedrigen pH-Werten kommt es zu verminderten Taxazahlen, allerdings sind viele Quellorganismen offen-sichtlich an eine leichte Versauerung angepasst, da in sauren Quellen der Anteil der Quellfauna höher war. Außerdem ergab die Untersuchung, dass neue und intakte Fassungen weniger Taxa und Quelltaxa aufwiesen, während bei alten und verfallenen Fassungen oft eine hohe Quelltaxazahl vorlag, so dass diese als sekundär ökologisch wertvoll zu bezeichnen sind. Die Umfeldnutzung und die Größe des Quellbereichs sind ebenfalls wichtig, so dass Quellen in Laub- und Mischwald sowie in extensivem Grünland am besten besiedelt waren. Die stärksten statistischen Zusammenhänge mit der faunistischen Besiedlung ergaben sich bei den Bewer-tungsergebnissen des Strukturverfahrens, was zeigt, dass das Verfahren realistisch bewertet. Multivariate statistische Methoden dienten der Ermittlung von Besiedlungsfaktoren und der Leitbildentwick-lung. Es ergab sich eine unterschiedliche Besiedlung bei verschiedener Hydrogeologie und Hydrochemie sowie bei verschiedenen Quelltypen. Allerdings waren die Ergebnisse nicht eindeutig, so dass die Trennung der Quelltypen nur teilweise gelang und die Quelltypen von den Grundwasserlandschaften überlagert waren. Die Faunenunterschiede verschiedener Regionen waren größer als die einer Region, so dass morphologische Quelltypen faunistisch nur innerhalb einer hydrogeologischen Einheit von Bedeutung sind. Es wurden regionale faunistische Leitarten für vier Grundwasserlandschaften genannt, wobei sich auf wenige hochstete Quellarten beschränkt wurde. Die Arbeit zeigt, dass faunistisch orientierte Leitbilder nur unter Schwierigkeiten anzugeben sind und vollständige Typuszönosen für größere Gebiete grundsätzlich in Frage zu stellen sind. Sehr häufige grundsätzliche Leitarten für Quellen silikatischer Mittelgebirge in Rheinland-Pfalz sind Crunoecia irrorata und Bythinella dunkeri.