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Institute
Streams are coupled with their riparian area. Emerging insects from streams can be an important prey in the riparian area. Such aquatic subsidies can cause predators to switch prey or increase predator abundances. This can impact the whole terrestrial food web. Stressors associated with agricultural land use can alter insect communities in water and on land, resulting in complex response patterns of terrestrial predators that rely on prey from both systems.
This thesis comprises studies on the impact of aquatic nsects on a terrestrial model ecosystem (Objective 1, hapter 2), the influence of agricultural land use on riparian spiders’ traits and community (Objective 2, Chapter 3), and on the impact of agricultural land use on the contribution of different prey to spider diet (Objective 3, Chapter 4).
In chapter 2, I present a study where we conducted a mesocosm experiment to examine the effects of aquatic subsidies on a simplified terrestrial food web consisting of two types of herbivores (leafhoppers and weevils), plants and predators (spiders). I focused on the prey choice of the spiders by excluding predator immigration and reproduction. In accordance with predator switching, survival of leafhoppers increased in the presence of aquatic subsidies. By contrast, the presence of aquatic subsidies indirectly reduced weevils and herbivory.
In chapter 3, I present the results on the taxonomic and trait response of riparian spider communities to gradients of agricultural stressors and environmental variables, with a particular emphasis on pesticides. To capture spiders with different traits and survival strategies, we used multiple collection methods. Spider community composition was best explained by in-stream pesticide toxicity and shading of the stream bank, a proxy for the quality of the habitat. Species richness and the number of spider individuals, as well as community ballooning ability, were negatively associated with in-stream pesticide toxicity. In contrast, mean body size and shading preference of spider communities responded strongest to shading,
whereas mean niche width (habitat preference for moisture and shading) responded strongest to other environmental variables.
In chapter 4, I describe aquatic-terrestrial predator-prey relations with gradients of agricultural stressors and environmental variables. I sampled spiders, as well as their aquatic and terrestrial prey along streams with an assumed pesticide pollution gradient and determined their stable carbon and nitrogen signals. Potential aquatic prey biomass correlated positively with an increasing aquatic prey contribution of T. montana. The contribution of aquatic prey to the diet of P. amentata showed a positive relationship with increasing toxicity in streams.
Overall, this thesis contributes to the emerging discipline of cross-ecosystem ecology and shows that aquatic-terrestrial linkages and riparian food webs can be influenced by land use related stressors. Future manipulative field studies on aquatic-terrestrial linkages are required that consider the quality of prey organisms, fostering mechanistic understanding of such crossecosystem effects. Knowledge on these linkages is important to improve understanding of consequences of anthropogenic stressors and to prevent further losses of ecosystems and their biodiversity.
The growing numbers of breeding rooks (Corvus frugilegus) in the city of Landau (Rhineland- Palatinate, Germany) increase the potential for conflict between rooks and humans, which is mainly associated with noise and faeces. Therefore, the aim of this work is a better understanding of the breeding tree selection of the rook in order to develop options for action and management in the future.
Part I of this thesis provides general background information on the rook and includes mapping of the rookeries in the Anterior Palatinate and South Palatinate including Landau in the year 2020. That mapping revealed that the number of rural colonies has decreased, while the number of urban colonies has increased in the study area in the last few years. In line with current literature, tree species and tree size were important criteria for breeding tree selection. However, the mapping showed that additional factors must be important as well.
Therefore, as rooks seem to often breed along traffic axes, Part II of this thesis examines how temperature, artificial light and noise, which are all linked to traffic axes, affect the breeding tree selection of the rook in the city of Landau. The following three hypotheses are developed: (1) manually selected breeding trees (Bm) have a warmer microclimate than manually selected non-breeding trees (Nm) or randomly selected non-breeding trees (Nr), (2) Bm are exposed to a higher light level than Nm or Nr and (3) Bm are exposed to a higher noise level than Nm or Nr. To test these hypotheses, 15 Bm, 13 Nm and 16 Nr are investigated.
The results show that Bm were exposed to more noise than both types of non-breeding trees (μBm, noise = 36.52481 dB, μNm, noise = 31.27229 dB, μNr, noise = 29.17417 dB) where the difference between Bm and Nr was significant. In addition, there was a tendency for Bm to be exposed to less light (μBm, light = 0.356 lx) than Nm (μNm, light = 0.4107692 lx) and significantly less light than Nr (μNr, light = 1.995 lx), while temperature did not differ between the groups (μBm, temp = 16.90549 °C, μNm, temp = 16.93118 °C, μNr, temp = 17.28639 °C).
This study shows for the first time that rooks prefer trees which are exposed to low light levels and high noise levels, i.e. more intense traffic noise, for breeding. It can only be speculated that the cause of this is lower enemy pressure at such sites. The fact that temperature does not seem to have any influence on breeding tree selection may be due to only small temperature differences at nest height, which might be compensated by breeding behaviour. Consequently, in the long term one management approach could be to divert traffic from inner-city areas, especially schools and hospitals, to bypasses. If tree genera suitable for rooks, such as plane trees, are planted along the bypasses, those sites could provide suitable alternative habitats to inner-city breeding locations, which become less attractive for breeding due to noise reduction. In the short term in addition to locally implemented repellent measures the most effective approach is to strengthen rook acceptance among the population. However, further research is needed to verify the results of this thesis and to gain further insights into rook breeding site selection in order to develop effective management measures.
Flowering habitats to enhance biodiversity and pest control services in agricultural landscapes
(2015)
Meeting growing demands for agricultural products requires management solutions that enhance food production, whilst minimizing negative environmental impacts. Conventional agricultural intensification jeopardizes farmland biodiversity and associated ecosystem services through excessive anthropogenic inputs and landscape simplification. Agri-environment schemes (AES) are commonly implemented to mitigate the adverse effects of conventional intensification on biodiversity. However the moderate success of such schemes thus far would strongly benefit from more explicit goals regarding ecosystem service provisioning. Providing key resources to beneficial organisms may improve their abundance, fitness, diversity and the ecosystem services they provide. With targeted habitat management, AES may synergistically enhance biodiversity and agricultural production and thus contribute to ecological intensification. We demonstrate that sown perennial wildflower strips, as implemented in current AES focusing on biodiversity conservation also benefit biological pest control in nearby crops (Chapter 2).
Comparing winter wheat fields adjacent to wildflower strips with fields without wildflower strips we found strongly reduced cereal leaf beetle (Oulema sp.) density and plant damage near wildflower strips. In addition, winter wheat yield was 10 % higher when fields adjoined wildflower strips. This confirms previous assumptions that wildflower strips, known for positive effects on farmland biodiversity, can also enhance ecosystem services such as pest control and the positive correlation of yield with flower abundance and diversity suggests that floral resources are key. Refining sown flower strips for enhanced service provision requires mechanistic understanding of how organisms benefit from floral resources. In climate chamber experiments investigating the impact of single and multiple flowering plant species on fitness components of three key arthropod natural enemies of aphids, we demonstrate that different natural enemies benefit differently from the offered resources (Chapter 3).
Some flower species were hereby more valuable to natural enemies than others overall. Additionally, the mixture with all flowers generally performed better than monocultures, yet with no transgressive overyielding. By explicitly tailoring flower strips to the requirements of key natural enemies of crop pests we aimed to maximise natural enemy mediated pest control in winter wheat (Chapter 4)and potato (Chapter 5) crops.
Respecting the manifold requirements of diverse natural enemies but not pests, in terms of temporal and spatial provisioning of floral, extra floral and structural resources, we designed targeted annual flower strips that can be included in crop rotation to support key arthropods at the place and time they are needed. Indeed, field experiments revealed that cereal leaf beetle density and plant damage in winter wheat can be reduced by 40 % to 61 % and aphid densities in potatoes even by 77 %, if a targeted flower strip is sown into the field. These effects were not restricted to the vicinity of flower strips and, in contrast to fields without flower strip, often prevented action thresholds from being reached. This suggests that targeted flower strips could replace insecticides. All adult natural enemies were enhanced inside targeted flower strips when compared to control strips. Yet, spillover to the field was restricted to key natural enemies such as ground beetles (winter wheat), hoverflies (potato) and lacewings (winter wheat and potato), suggesting their dominant role in biological control. In potatoes, targeted flower strips also enhanced hoverfly species richness in strips and crop, highlighting their additional benefits for diversity.
The present results provide more insights into the mechanisms underlying conservation biological control and highlight the potential of tailored habitat management for ecological intensification.
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.
Wild bees are essential for the pollination of wild and cultivated plants. However, within the
last decades, the increasing intensification of modern agriculture has led to both a reduction and fragmentation as well as a degradation of the habitats wild bees need. The resulting loss of pollinators and their pollination poses an immense challenge to global food production. To support wild bees, the availability of flowering resources is essential. However, the flowering period of each resource is temporally limited and has different effects on pollinators and their pollination, depending on the time of their flowering.
Therefore, to efficiently promote and manage wild bee pollinators in agricultural landscapes, we identified species-specific key floral resources of three selected wild bee species and their spatial and temporal availability (CHAPTERS 2, 3 & 4). We examined, which habitat types predominantly provide these resources (CHAPTERS 3 & 4). We also investigated whether floral resource maps based on the use of these key resources and their spatial and temporal availability explain the abundance and development of the selected wild bees (CHAPTERS 3 & 4) and pollination (CHAPTER 5) better than habitat maps, that only indirectly account for the availability of floral resources.
For each of the species studied, we were able to identify different key pollen sources, predominantly woody plants in the early season (April/May) and increasingly herbaceous plants in the later season (June/July; CHAPTERS 2, 3 & 4). The open woody semi-natural habitats of our agricultural landscapes provided about 75% of the floral resources for the buff-tailed bumblebees, 60% for the red mason bees, and 55% for the horned mason bees studied, although they accounted for only 3% of the area (CHAPTERS 3 & 4). In addition, fruit orchards provided about 35% of the floral resources for the horned mason bees on 4% of the landscape area (CHAPTER 3). We showed that both mason bee species benefited from the resource availability in the surrounding landscapes (CHAPTER 3). Yet this was not the case for the bumblebees (CHAPTER 4). Instead, the weight gain of their colonies, the number of developed queen cells and their colony survival were higher with increasing proximity to forests. The proximity to forests also had a positive effect on the mason bees studied (CHAPTER 3). In addition, the red mason bees benefited from herbaceous semi-natural habitats. The proportion of built-up areas had a negative effect on the horned mason bees, and the proportion of arable land on the red mason bees. The habitat maps explained horned mason bee abundances equally well as the floral resource maps, but red mason bee abundances were distinctly better explained by key floral resources. The pollination of field bean increased with higher proportions of early floral resources, whereas synchronous floral resources showed no measurable reduction in their pollination (CHAPTER 5). Habitat maps also explained field bean pollination better than floral resource maps. Here, pollination increased with increasing proportions of built-up areas in the landscapes and decreased with increasing proportions of arable land.
Our results highlight the importance of the spatio-temporal availability of certain key species as resource plants of wild bees in agricultural landscapes. They show that habitat maps are ahead of, or at least equal to, spatio-temporally resolved floral resource maps in predicting wild bee development and pollination. Nevertheless, floral resource maps allow us to draw more accurate conclusions between key floral resources and the organisms studied. The proximity to forest edges had a positive effect on each of the three wild bee species studied. However, besides pure food availability, other factors seem to co-determine the occurrence of wild bees in agricultural landscapes.
The role of alternative resources for pollinators and aphid predators in agricultural landscapes
(2021)
The world wide decline of insects is often associated with loss of natural and semi-natural habitat caused by intensified land-use. Many insects provide important ecosystem services to agriculture, such as pest control or pollination. To efficiently promote insects on remaining semi-natural habitat we need precise knowledge of their requirements to non-crop habitat. This thesis focuses on identifying
the most important semi-natural habitats (forest edges, grasslands, and semi-open habitats) for pollinators and natural enemies of crop pests with respect to their food resource requirements. Special
attention is given to floral resources and their spatio-temporal distribution in agricultural landscapes.
Floral resource maps might get closer at characterizing landscapes the way they are experienced by insects compared to classical habitat maps. Performance of the two map types was compared on the prediction of wild bees and natural enemies that consume nectar and pollen, identifying habitats of special importance in the process. In wild bees, influences of spatio-temporal floral resource availability were analysed as well as habitat preferences of specific groups of bees. Understanding dietary needs of natural enemies of crop pests requires additional knowledge on prey use. To this end, ladybird gut contents have been analysed by means of high-throughput sequencing for insight into aphid prey-use.
Results showed, that wild bees were predicted better by floral resource maps compared to classical habitat maps. Forest edge area, as well as floral resources in forest edges had positive effects on abundance and diversity of rare bees and important crop pollinators. Similar patterns were retained for grassland diversity. Especially early floral resources seemed to have positive effects on wild bees. Crops and fruit trees produced a resource pulse in April that exceeded floral resource availability in May and June by tenfold. Most floral resources in forest edges appeared early in the season, with the highest floral density per area. Grasslands provided the lowest amount of floral resources but highest diversity, which was evenly distributed over the season.
Despite natural enemies need for floral resources, classical habitat maps performed better at predicting natural enemies of crop pests compared to floral resource maps. Classical habitat maps revealed a positive effect of forest edge habitat on the abundance of pest enemies, which translated into improved aphid control. Results from gut content analysis reveal high portions of pest aphid species and nettle aphids as well as a broader insight into prey spectra retained from ladybirds collected from sticky traps compared to individuals collected by hand. The aphid specific primer designed for this purpose will be helpful for identifying aphid consumption by ladybirds in future studies.
Findings of this thesis show the potential of floral resource maps for understanding interactions of wild bees and the landscape but also indicate that natural enemies are limited by other resources. I would like to highlight the positive effects of forest edges for different groups of bees as well as natural enemies and their performance on pest control.
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.
Invasive species play increasing roles worldwide. Invasions are considered successful when species establish and spread in their exotic range. Subsequently, dispersal is a major determinant of species’ range dynamics. Mermessus trilobatus, native to North America, has rapidly spread in Europe via aerial dispersal. Here we investigated the interplay of ecological and evolutionary processes behind its colonisation success.
First, we examined two possible ecological mechanisms. Similar to other invasive invertebrates, the colonisation success of Mermessus trilobatus might be related to human-induced habitat disturbance. Opposite to this expectation, our results showed that densities of Mermessus trilobatus decreased with soil disturbance in grasslands suggesting that its invasion success was not connected to a ruderal strategy. Further, invasive species often escape the ecological pressures from novel enemies in their exotic ranges. Unexpectedly, invasive Mermessus trilobatus was more sensitive to a native predator than native Erigone dentipalpis during our predator susceptibility trials. This indicates that the relation between the invasive spider and its native predator is dominated by prey naïveté rather than enemy release.
The remaining three chapters of the thesis investigated the dispersal behaviour of this invasive species. Hitherto, studies of passive aerial dispersal used wind as the primary dispersal-initiating factor despite a recent demonstration of the effects of the atmospheric electric fields on spiders’ pre-dispersal behaviour. During our experiments, only the wind facilitated the flight, although electric fields induced pre-dispersal behaviour in spiders. Consequently, studies around passive aerial dispersal should control electric fields but use wind as a stimulating factor.
Rapidly expanding species might be disproportionately distributed in their exotic range, with an accumulation of dispersive genotypes at the leading edge of their range. Such imbalanced spatial segregation is possible when the dispersal behaviour of expanding species is heritable. Our results showed that the dispersal traits of Mermessus trilobatus were heritable through both parents and for both sexes with recessive inheritance of high dispersal ability in this species.
Following the heritability experiments, we documented an accelerated spread of Mermessus trilobatus in Europe and tested whether dispersal, reproduction or competing ability was at the source of this pattern. Our results showed that the accumulation of more mobile but not reproductive or competitive genotypes at the expansion front of this invasive species gave rise to an accelerated range expansion by more than 1350 km in under 45 years.
Invasive Mermessus trilobatus is inferior to native sympatric species with respect to competing ability (Eichenberger et al., 2009), disturbance tolerance and predation pressure. Nevertheless, the species successfully established in its exotic range and spread by accelerating its expansion rate. Rapid reproduction that balances the high ecological pressures might be the other potential mechanism behind its colonisation success in Europe and deserves further investigation.
Non-Consumptive Effects of Spiders and Ants: Does Fear Matter in Terrestrial Interaction Webs?
(2014)
Most animals suffer from predators. Besides killing prey, predators can affect prey physiology, morphology and behaviour. Spiders are among the most diverse and frequent predators in terrestrial ecosystems. Our behavioural arena experiments revealed that behavioural changes under spider predation risk are relatively scarce among arthropods. Wood crickets (Nemobius sylvestris), in particular, changed their behaviour in response to cues of various spider species. Thereby, more common and relatively larger spider species induced stronger antipredator behaviour in crickets.
Behavioural changes under predation risk are expected to enhance predator avoidance, but they come at a cost. Crickets previously confronted with cues of the nursery web spider (Pisaura mirabilis) were indeed more successful in avoiding predation. Surprisingly, crickets slightly increased food uptake and lost less weight under predation risk, indicating that crickets are able to compensate for short-term cost under predation risk. In a following plant choice experiment, crickets strongly avoided plants bearing spider cues, which in turn reduced the herbivory on the respective plants.
Similar to spiders, ants are ubiquitous predators and can have a strong impact on herbivores, but also on other predators. Juvenile spiders increased their propensity for long-distance dispersal if exposed to ant cues. Thus, spiders use this passive dispersal through the air (ballooning) to avoid ants and colonise new habitats.
In a field experiment, we compared arthropod colonisation between plants bearing cues of the nursery web spider and cue-free plants. We followed herbivory during the experimental period and sampled the arthropod community on the plants. In accordance with the plant choice experiment, herbivory was reduced on plants bearing spider cues. In addition, spider cues led to changes in the arthropod community: smaller spiders and black garden ants (Lasius niger) avoided plants bearing spider cues. In contrast, common red ants (Myrmica rubra) increased the recruitment of workers, possibly to protect their aphids.
Although behavioural changes were relatively rare on filter papers bearing spider cues, more natural experimental setups revealed strong and far-reaching effects of predation risk. We further suggest that risk effects influence the spatial distribution of herbivory, rather than reduce overall herbivory that is expected if predators kill herbivores. Consequently, the relative importance of predation and risk effects is crucial for the way predators affect lower trophic levels.
Carabids, which are frequently distributed in agricultural landscapes, are natural enemies of different pests including slugs. Semi-natural habitats are known to affect carabids and thus, their potential to support natural pest control.
The impact of semi-natural habitats was investigated on carabids and slugs within different non-crop habitats (chapter 2). Most carabids and Deroceras reticulatum showed preferences for herbaceous semi-natural habitats, while Arion spp. occured mainly in woody habitats. An increase of predatory carabid abundance, which was linked to an inclining amount of semi-natural habitats in the landscape, and a decrease of Arion spp. densities, indicated a high potential for slug control in structural rich landscapes.
Effects of semi-natural habitats were investigated on predatory carabids and slugs in 18 wheat fields (chapter 3). Predatory carabid species richness was positively affected by the increasing amount of semi-natural habitats in the landscape, whereas predatory carabid abundance was neither influenced by adjacent habitat type nor by the proportion of semi-natural habitats in the landscape. The target pest species showed divergent patterns, whereas Arion spp. densities were highest in structural poor landscapes near woody margins. D. reticulatum was not affected by habitat type or landscape, reflecting its adaptation to agriculture. Results indicate an increased control of Arion spp. by carabids in landscapes with a high amount of semi-natural habitats.
Effects of semi-natural habitats and the influence of farming system was tested on carabid distribution within 18 pumpkin fields (chapter 4). Carabid species richness generally increased with decreasing distance to the field margins, whereas carabid abundance responded differently according to the adjacent habitat type. Farming system had no effect on carabids and landscape heterogeneity only affected carabids in organic pumpkin fields.
Slug and slug egg predation of three common carabid species was tested in single and double species treatments in the laboratory (chapter 5). Results show additive and synergistic effects depending on the carabid species. In general, semi-natural habitats can enhance the potential of slug control by carabids. This counts especially for Arionid slugs. Semi-natural habitats can support carabid communities by providing shelter, oviposition and overwintering sites as wells as complementary food sources. Therefore, it is important to provide a certain amount of non-crop habitats in agricultural landscapes.