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INTRODUCTION: The proximity of semi-natural habitats and agricultural fields in an agricultural landscape leads to unavoidable biological, chemical, and physical interactions. Fungi can negatively influence, but also support crop growth in agricultural fields. Therefore, in this field study we investigated the colonisation of arbuscular mycorrhizal (AM) fungi and non-AM fungi in winter-wheat roots as well as winter-wheat performance in distance to semi-natural habitats. MATERIALS AND METHODS: We sampled in an intensively managed agricultural landscape in North-east Germany along agricultural transition zones, that is, along 50 m-transects from semi-natural habitats like hedgerows and glacially created in-field ponds—so-called kettle holes—into agricultural fields. RESULTS: To our knowledge, we show for the first time that AM fungal colonisation in winter-wheat roots decreased linearly with increasing distance to semi-natural habitats while non-AM fungal root colonisation did not change. Winter-wheat grain yield and biomass slightly increased with increasing distance to hedgerows but not to kettle holes. This clearly shows that there is a difference between different crop performance parameters. Random forest machine learning algorithms confirmed the particular importance of distance to semi-natural habitats for AM fungal root colonisation and for winter-wheat grain yield. Less intensive agricultural management close to semi-natural habitats, for example, no herbicide and pesticide applications as a result of nature protection regulations, may partly explain this pattern. However, spatial response patterns of AM but not of non-AM fungi in wheat roots also point to changed ecological interactions close to semi-natural habitats. CONCLUSION: Semi-natural and natural habitats in agricultural landscapes are slowly recognised not only to be important for biodiversity conservation, but also for sustainable crop production. Additionally, they may also be a tool for farmers and policy makers to improve sustainable landscape management. And agricultural transition zones are spatially and temporally complex dynamic ecosystems that should be the focus of further investigations.

Semi-natural habitats (SNHs) in agricultural landscapes represent important refugia for biodiversity including organisms providing ecosystem services. Their spill-over into agricultural fields may lead to the provision of regulating ecosystem services such as biological pest control ultimately affecting agricultural yield. Still, it remains largely unexplored, how different habitat types and their distributions in the surrounding landscape shape this provision of ecosystem services within arable fields. Hence, in this thesis I investigated the effect of SNHs on biodiversity-driven ecosystem services and disservices affecting wheat production with an emphasis on the role and interplay of habitat type, distance to the habitat and landscape complexity. I established transects from the field border into the wheat field, starting either from a field-to-field border, a hedgerow, or a kettle hole, and assessed beneficial and detrimental organisms and their ecosystem functions as well as wheat yield at several in-field distances. Using this study design, I conducted three studies where I aimed to relate the impacts of SNHs at the field and at the landscape scale on ecosystem service providers to crop production. In the first study, I observed yield losses close to SNHs for all transect types. Woody habitats, such as hedgerows, reduced yields stronger than kettle holes, most likely due to shading from the tall vegetation structure. In order to find the biotic drivers of these yield losses close to SNHs, I measured pest infestation by selected wheat pests as potential ecosystem disservices to crop production in the second study. Besides relating their damage rates to wheat yield of experimental plots, I studied the effect of SNHs on these pest rates at the field and at the landscape scale. Only weed cover could be associated to yield losses, having their strongest impact on wheat yield close to the SNH. While fungal seed infection rates did not respond to SNHs, fungal leaf infection and herbivory rates of cereal leaf beetle larvae were positively influenced by kettle holes. The latter even increased at kettle holes with increasing landscape complexity suggesting a release of natural enemies at isolated habitats within the field interior. In the third study, I found that also ecosystem service providers benefit from the presence of kettle holes. The distance to a SNH decreased species richness of ecosystem service providers, whereby the spatial range depended on species mobility, i.e. arable weeds diminished rapidly while carabids were less affected by the distance to a SNH. Contrarily, weed seed predation increased with distance suggesting that a higher food availability at field borders might have diluted the predation on experimental seeds. Intriguingly, responses to landscape complexity were rather mixed: While weed species richness was generally elevated with increasing landscape complexity, carabids followed a hump-shaped curve with highest species numbers and activity-density in simple landscapes. The latter might give a hint that carabids profit from a minimum endowment of SNHs, while a further increase impedes their mobility. Weed seed predation was affected differently by landscape complexity depending on weed species displayed. However, in habitat-rich landscapes seed predation of the different weed species converged to similar rates, emphasising that landscape complexity can stabilize the provision of ecosystem services. Lastly, I could relate a higher weed seed predation to an increase in wheat yield even though seed predation did not diminish weed cover. The exact mechanisms of the provision of weed control to crop production remain to be investigated in future studies. In conclusion, I found habitat-specific responses of ecosystem (dis)service providers and their functions emphasizing the need to evaluate the effect of different habitat types on the provision of ecosystem services not only at the field scale, but also at the landscape scale. My findings confirm that besides identifying species richness of ecosystem (dis)service providers the assessment of their functions is indispensable to relate the actual delivery of ecosystem (dis)services to crop production. Naturnahe Habitate, wie zum Beispiel Hecken und Sölle, stellen wichtige Refugien für die Biodiversität in Agrarlandschaften dar, weil aus diesen Habitaten Organismen in die Agrarflächen einwandern und dort regulierende Ökosystemdienstleistungen, wie zum Beispiel biologische Schädlingsbekämpfung, erbringen können. Weitgehend unerforscht ist bisher, in welcher Art und Weise die verschiedenen Habitattypen und ihre Verteilung in der umgebenden Landschaft die Bereitstellung dieser Ökosystemdienstleistungen, die letztlich auch einen Einfluss auf die landwirtschaftlichen Erträge haben können, beeinflussen. Daher habe ich den Einfluss von naturnahen Habitattypen auf biodiversitätsbedingte Ökosystemdienstleistungen und ihre Auswirkung auf die Weizenproduktion untersucht. Der Schwerpunkt meiner Arbeit lag auf dem Einfluss und dem Zusammenspiel von Habitattyp, Entfernung zum naturnahen Habitat und der umgebenden Landschaftsvielfalt. Auf intensiv bewirtschafteten Weizenfeldern habe ich entlang von Transekten von der Feldgrenze in das Feld hinein Nützlinge und Schädlinge, Ökosystemfunktionen sowie den Weizenertrag ermittelt. In der ersten Studie habe ich am Feldrand für alle Habitattypen einen Ertragsverlust im Vergleich zur Feldmitte beobachtet, wobei Hecken die stärkste Ertragsreduktion aufwiesen. Dieses Resultat führe ich auf die Beschattung durch die hohe Vegetationsstruktur zurück. Um Ertragsverluste besser zu verstehen, habe ich in der zweiten Studie den Schädlingsbefall durch ausgewählte Weizenschädlinge sowie den direkten Einfluss der naturnahen Habitate auf die Schädlingsraten auf der Feld- und Landschaftsskala untersucht. Nur die Unkrautbedeckung konnte mit dem Ertragsverlust in Verbindung gebracht werden, wobei sie einen stärkeren Einfluss auf die Ernteerträge in der Nähe der naturnahen Habitate hatte. Darüber hinaus konnte ich zeigen, dass die Befallsraten von Blattpathogenen und die Fraßraten der Larven des Getreidehähnchens an Söllen erhöht waren. Letzteres stieg sogar an Söllen mit zunehmender Landschaftsvielfalt an, was auf den Wegfall natürlicher Feinde an isolierten Habitaten im Feldinneren, wie Söllen, schließen lässt. In meiner dritten Studie fand ich heraus, dass auch Ökosystemdienstleister, wie Laufkäfer sowie die Samenprädation von Unkräutern, von Söllen profitieren. Die Entfernung zu einem naturnahen Habitat verringerte den Artenreichtum der Ökosystemdienstleister, im Gegensatz zur Samenprädation, welche zur Feldmitte zunahm. Dies deutet darauf hin, dass eine höhere Nahrungsverfügbarkeit an Feldrändern die Prädation von Versuchssamen abgeschwächt haben könnte. Während mit zunehmender Landschaftsvielfalt die Artenanzahl an Unkräutern anstieg, war bei den Laufkäfern die höchste Artenzahl und Aktivitätsdichte in Landschaften mit geringer Vielfalt zu beobachten. Das lässt den Schluss zu, dass eine Minimalausstattung an naturnahen Habitaten für Laufkäfer vorteilhaft ist, während ein zu großer Anteil an naturnahen Habitaten ihre Mobilität behindern könnte. Die ausgelegten Samen wurden in habitatarmen Landschaften unterschiedlich stark gefressen, wohingegen sie sich in habitatreichen Landschaften anglichen. Dieses Resultat unterstreicht, dass Landschaftsvielfalt die Bereitstellung von Ökosystemdienstleistungen stabilisieren kann. Abschließend konnte ich zeigen, dass mit ansteigender Samenprädation von Unkräutern ein Anstieg des Weizenertrages einherging, wenn auch die Unkrautbedeckung nicht verringert wurde. Die genauen Mechanismen der Bereitstellung von natürlicher Unkrautbekämpfung für die Pflanzenproduktion sollten in zukünftigen Studien weiter untersucht werden. Zusammenfassend lässt sich sagen, dass die untersuchten Ökosystemdienstleister und ihre Schädlings- sowie Prädationsraten Habitatpräferenzen aufwiesen. Diese Tatsache unterstreicht die Notwendigkeit, die Auswirkungen verschiedener Habitattypen auf die Bereitstellung von Ökosystemdienstleistungen nicht nur auf der Feldskala, sondern auch auf der Landschaftsskala zu bewerten. Meine Ergebnisse bestätigen, dass neben der Aufnahme des Artenreichtums von Ökosystemdienstleistern die Bewertung ihrer Funktionen unerlässlich ist, um die tatsächliche Bereitstellung von Ökosystemdienstleistungen mit dem landwirtschaftlichen Ertrag in Beziehung zu setzen.

Semi‐natural habitats (SNHs) are becoming increasingly scarce in modern agricultural landscapes. This may reduce natural ecosystem services such as pest control with its putatively positive effect on crop production. In agreement with other studies, we recently reported wheat yield reductions at field borders which were linked to the type of SNH and the distance to the border. In this experimental landscape‐wide study, we asked whether these yield losses have a biotic origin while analyzing fungal seed and fungal leaf pathogens, herbivory of cereal leaf beetles, and weed cover as hypothesized mediators between SNHs and yield. We established experimental winter wheat plots of a single variety within conventionally managed wheat fields at fixed distances either to a hedgerow or to an in‐field kettle hole. For each plot, we recorded the fungal infection rate on seeds, fungal infection and herbivory rates on leaves, and weed cover. Using several generalized linear mixed‐effects models as well as a structural equation model, we tested the effects of SNHs at a field scale (SNH type and distance to SNH) and at a landscape scale (percentage and diversity of SNHs within a 1000‐m radius). In the dry year of 2016, we detected one putative biotic culprit: Weed cover was negatively associated with yield values at a 1‐m and 5‐m distance from the field border with a SNH. None of the fungal and insect pests, however, significantly affected yield, neither solely nor depending on type of or distance to a SNH. However, the pest groups themselves responded differently to SNH at the field scale and at the landscape scale. Our findings highlight that crop losses at field borders may be caused by biotic culprits; however, their negative impact seems weak and is putatively reduced by conventional farming practices., Understanding how semi‐natural habitats influence provisioning ecosystem services of agricultural fields should be a key argument in debates about economic profits versus biodiversity conservation in agricultural landscapes. Therefore, we studied the effect of pest infestation as a biotic culprit to yield losses close to hedgerows and kettle holes using a structural equation model. We found arable weed cover being associated with yield losses along the transects, however, only in the proximate distances to the field border.

Semi-natural habitats (SNHs) in agricultural landscapes represent important refugia for biodiversity including organisms providing ecosystem services. Their spill-over into agricultural fields may lead to the provision of regulating ecosystem services such as biological pest control ultimately affecting agricultural yield. Still, it remains largely unexplored, how different habitat types and their distributions in the surrounding landscape shape this provision of ecosystem services within arable fields. Hence, in this thesis I investigated the effect of SNHs on biodiversity-driven ecosystem services and disservices affecting wheat production with an emphasis on the role and interplay of habitat type, distance to the habitat and landscape complexity. I established transects from the field border into the wheat field, starting either from a field-to-field border, a hedgerow, or a kettle hole, and assessed beneficial and detrimental organisms and their ecosystem functions as well as wheat yield at several in-field distances. Using this study design, I conducted three studies where I aimed to relate the impacts of SNHs at the field and at the landscape scale on ecosystem service providers to crop production. In the first study, I observed yield losses close to SNHs for all transect types. Woody habitats, such as hedgerows, reduced yields stronger than kettle holes, most likely due to shading from the tall vegetation structure. In order to find the biotic drivers of these yield losses close to SNHs, I measured pest infestation by selected wheat pests as potential ecosystem disservices to crop production in the second study. Besides relating their damage rates to wheat yield of experimental plots, I studied the effect of SNHs on these pest rates at the field and at the landscape scale. Only weed cover could be associated to yield losses, having their strongest impact on wheat yield close to the SNH. While fungal seed infection rates did not respond to SNHs, fungal leaf infection and herbivory rates of cereal leaf beetle larvae were positively influenced by kettle holes. The latter even increased at kettle holes with increasing landscape complexity suggesting a release of natural enemies at isolated habitats within the field interior. In the third study, I found that also ecosystem service providers benefit from the presence of kettle holes. The distance to a SNH decreased species richness of ecosystem service providers, whereby the spatial range depended on species mobility, i.e. arable weeds diminished rapidly while carabids were less affected by the distance to a SNH. Contrarily, weed seed predation increased with distance suggesting that a higher food availability at field borders might have diluted the predation on experimental seeds. Intriguingly, responses to landscape complexity were rather mixed: While weed species richness was generally elevated with increasing landscape complexity, carabids followed a hump-shaped curve with highest species numbers and activity-density in simple landscapes. The latter might give a hint that carabids profit from a minimum endowment of SNHs, while a further increase impedes their mobility. Weed seed predation was affected differently by landscape complexity depending on weed species displayed. However, in habitat-rich landscapes seed predation of the different weed species converged to similar rates, emphasising that landscape complexity can stabilize the provision of ecosystem services. Lastly, I could relate a higher weed seed predation to an increase in wheat yield even though seed predation did not diminish weed cover. The exact mechanisms of the provision of weed control to crop production remain to be investigated in future studies. In conclusion, I found habitat-specific responses of ecosystem (dis)service providers and their functions emphasizing the need to evaluate the effect of different habitat types on the provision of ecosystem services not only at the field scale, but also at the landscape scale. My findings confirm that besides identifying species richness of ecosystem (dis)service providers the assessment of their functions is indispensable to relate the actual delivery of ecosystem (dis)services to crop production., Naturnahe Habitate, wie zum Beispiel Hecken und Sölle, stellen wichtige Refugien für die Biodiversität in Agrarlandschaften dar, weil aus diesen Habitaten Organismen in die Agrarflächen einwandern und dort regulierende Ökosystemdienstleistungen, wie zum Beispiel biologische Schädlingsbekämpfung, erbringen können. Weitgehend unerforscht ist bisher, in welcher Art und Weise die verschiedenen Habitattypen und ihre Verteilung in der umgebenden Landschaft die Bereitstellung dieser Ökosystemdienstleistungen, die letztlich auch einen Einfluss auf die landwirtschaftlichen Erträge haben können, beeinflussen. Daher habe ich den Einfluss von naturnahen Habitattypen auf biodiversitätsbedingte Ökosystemdienstleistungen und ihre Auswirkung auf die Weizenproduktion untersucht. Der Schwerpunkt meiner Arbeit lag auf dem Einfluss und dem Zusammenspiel von Habitattyp, Entfernung zum naturnahen Habitat und der umgebenden Landschaftsvielfalt. Auf intensiv bewirtschafteten Weizenfeldern habe ich entlang von Transekten von der Feldgrenze in das Feld hinein Nützlinge und Schädlinge, Ökosystemfunktionen sowie den Weizenertrag ermittelt. In der ersten Studie habe ich am Feldrand für alle Habitattypen einen Ertragsverlust im Vergleich zur Feldmitte beobachtet, wobei Hecken die stärkste Ertragsreduktion aufwiesen. Dieses Resultat führe ich auf die Beschattung durch die hohe Vegetationsstruktur zurück. Um Ertragsverluste besser zu verstehen, habe ich in der zweiten Studie den Schädlingsbefall durch ausgewählte Weizenschädlinge sowie den direkten Einfluss der naturnahen Habitate auf die Schädlingsraten auf der Feld- und Landschaftsskala untersucht. Nur die Unkrautbedeckung konnte mit dem Ertragsverlust in Verbindung gebracht werden, wobei sie einen stärkeren Einfluss auf die Ernteerträge in der Nähe der naturnahen Habitate hatte. Darüber hinaus konnte ich zeigen, dass die Befallsraten von Blattpathogenen und die Fraßraten der Larven des Getreidehähnchens an Söllen erhöht waren. Letzteres stieg sogar an Söllen mit zunehmender Landschaftsvielfalt an, was auf den Wegfall natürlicher Feinde an isolierten Habitaten im Feldinneren, wie Söllen, schließen lässt. In meiner dritten Studie fand ich heraus, dass auch Ökosystemdienstleister, wie Laufkäfer sowie die Samenprädation von Unkräutern, von Söllen profitieren. Die Entfernung zu einem naturnahen Habitat verringerte den Artenreichtum der Ökosystemdienstleister, im Gegensatz zur Samenprädation, welche zur Feldmitte zunahm. Dies deutet darauf hin, dass eine höhere Nahrungsverfügbarkeit an Feldrändern die Prädation von Versuchssamen abgeschwächt haben könnte. Während mit zunehmender Landschaftsvielfalt die Artenanzahl an Unkräutern anstieg, war bei den Laufkäfern die höchste Artenzahl und Aktivitätsdichte in Landschaften mit geringer Vielfalt zu beobachten. Das lässt den Schluss zu, dass eine Minimalausstattung an naturnahen Habitaten für Laufkäfer vorteilhaft ist, während ein zu großer Anteil an naturnahen Habitaten ihre Mobilität behindern könnte. Die ausgelegten Samen wurden in habitatarmen Landschaften unterschiedlich stark gefressen, wohingegen sie sich in habitatreichen Landschaften anglichen. Dieses Resultat unterstreicht, dass Landschaftsvielfalt die Bereitstellung von Ökosystemdienstleistungen stabilisieren kann. Abschließend konnte ich zeigen, dass mit ansteigender Samenprädation von Unkräutern ein Anstieg des Weizenertrages einherging, wenn auch die Unkrautbedeckung nicht verringert wurde. Die genauen Mechanismen der Bereitstellung von natürlicher Unkrautbekämpfung für die Pflanzenproduktion sollten in zukünftigen Studien weiter untersucht werden. Zusammenfassend lässt sich sagen, dass die untersuchten Ökosystemdienstleister und ihre Schädlings- sowie Prädationsraten Habitatpräferenzen aufwiesen. Diese Tatsache unterstreicht die Notwendigkeit, die Auswirkungen verschiedener Habitattypen auf die Bereitstellung von Ökosystemdienstleistungen nicht nur auf der Feldskala, sondern auch auf der Landschaftsskala zu bewerten. Meine Ergebnisse bestätigen, dass neben der Aufnahme des Artenreichtums von Ökosystemdienstleistern die Bewertung ihrer Funktionen unerlässlich ist, um die tatsächliche Bereitstellung von Ökosystemdienstleistungen mit dem landwirtschaftlichen Ertrag in Beziehung zu setzen.

Natural landscape elements (NLEs) in agricultural landscapes contribute to biodiversity and ecosystem services, but are also regarded as an obstacle for large‐scale agricultural production. However, the effects of NLEs on crop yield have rarely been measured. Here, we investigated how different bordering structures, such as agricultural roads, field‐to‐field borders, forests, hedgerows, and kettle holes, influence agricultural yields. We hypothesized that (a) yield values at field borders differ from mid‐field yields and that (b) the extent of this change in yields depends on the bordering structure. We measured winter wheat yields along transects with log‐scaled distances from the border into the agricultural field within two intensively managed agricultural landscapes in Germany (2014 near Göttingen, and 2015–2017 in the Uckermark). We observed a yield loss adjacent to every investigated bordering structure of 11%–38% in comparison with mid‐field yields. However, depending on the bordering structure, this yield loss disappeared at different distances. While the proximity of kettle holes did not affect yields more than neighboring agricultural fields, woody landscape elements had strong effects on winter wheat yields. Notably, 95% of mid‐field yields could already be reached at a distance of 11.3 m from a kettle hole and at a distance of 17.8 m from hedgerows as well as forest borders. Our findings suggest that yield losses are especially relevant directly adjacent to woody landscape elements, but not adjacent to in‐field water bodies. This highlights the potential to simultaneously counteract yield losses close to the field border and enhance biodiversity by combining different NLEs in agricultural landscapes such as creating strips of extensive grassland vegetation between woody landscape elements and agricultural fields. In conclusion, our results can be used to quantify ecocompensations to find optimal solutions for the delivery of productive and regulative ecosystem services in heterogeneous agricultural landscapes., Postprints der Universität Potsdam : Mathematisch-Naturwissenschaftliche Reihe, 811

Natural landscape elements (NLEs) in agricultural landscapes contribute to biodiversity and ecosystem services, but are also regarded as an obstacle for large‐scale agricultural production. However, the effects of NLEs on crop yield have rarely been measured. Here, we investigated how different bordering structures, such as agricultural roads, field‐to‐field borders, forests, hedgerows, and kettle holes, influence agricultural yields. We hypothesized that (a) yield values at field borders differ from mid‐field yields and that (b) the extent of this change in yields depends on the bordering structure. We measured winter wheat yields along transects with log‐scaled distances from the border into the agricultural field within two intensively managed agricultural landscapes in Germany (2014 near Göttingen, and 2015–2017 in the Uckermark). We observed a yield loss adjacent to every investigated bordering structure of 11%–38% in comparison with mid‐field yields. However, depending on the bordering structure, this yield loss disappeared at different distances. While the proximity of kettle holes did not affect yields more than neighboring agricultural fields, woody landscape elements had strong effects on winter wheat yields. Notably, 95% of mid‐field yields could already be reached at a distance of 11.3 m from a kettle hole and at a distance of 17.8 m from hedgerows as well as forest borders. Our findings suggest that yield losses are especially relevant directly adjacent to woody landscape elements, but not adjacent to in‐field water bodies. This highlights the potential to simultaneously counteract yield losses close to the field border and enhance biodiversity by combining different NLEs in agricultural landscapes such as creating strips of extensive grassland vegetation between woody landscape elements and agricultural fields. In conclusion, our results can be used to quantify ecocompensations to find optimal solutions for the delivery of productive and regulative ecosystem services in heterogeneous agricultural landscapes.

Natural landscape elements (NLEs) in agricultural landscapes contribute to biodiversity and ecosystem services, but are also regarded as an obstacle for large‐scale agricultural production. However, the effects of NLEs on crop yield have rarely been measured. Here, we investigated how different bordering structures, such as agricultural roads, field‐to‐field borders, forests, hedgerows, and kettle holes, influence agricultural yields. We hypothesized that (a) yield values at field borders differ from mid‐field yields and that (b) the extent of this change in yields depends on the bordering structure. We measured winter wheat yields along transects with log‐scaled distances from the border into the agricultural field within two intensively managed agricultural landscapes in Germany (2014 near Göttingen, and 2015–2017 in the Uckermark). We observed a yield loss adjacent to every investigated bordering structure of 11%–38% in comparison with mid‐field yields. However, depending on the bordering structure, this yield loss disappeared at different distances. While the proximity of kettle holes did not affect yields more than neighboring agricultural fields, woody landscape elements had strong effects on winter wheat yields. Notably, 95% of mid‐field yields could already be reached at a distance of 11.3 m from a kettle hole and at a distance of 17.8 m from hedgerows as well as forest borders. Our findings suggest that yield losses are especially relevant directly adjacent to woody landscape elements, but not adjacent to in‐field water bodies. This highlights the potential to simultaneously counteract yield losses close to the field border and enhance biodiversity by combining different NLEs in agricultural landscapes such as creating strips of extensive grassland vegetation between woody landscape elements and agricultural fields. In conclusion, our results can be used to quantify ecocompensations to find optimal solutions for the delivery of productive and regulative ecosystem services in heterogeneous agricultural landscapes.