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Summary
Chapters I & II: General Introduction & General Methods
Agriculture is confronted with a rampant loss of biodiversity potentially eroding ecosystem service potentials and adding up to other stressors like climate change or the consequences of land-use change and intensive management. To counter this ‘biodiversity crisis’, agri-environment schemes (AES) have been introduced as part of ecological intensification efforts. These AES combine special management regimes with the establishment of tailored habitats to create refuges for biodiversity in agricultural landscapes and thus ensure biodiversity mediated ecosystem services such as pest control. However, little is known about how well different AES habitats fulfil this purpose and whether they benefit ecosystem services in adjacent crop fields. Here I investigated how effective different AES habitats are for restoring biodiversity in different agricultural landscapes (Chapter V) and whether they benefit natural pest control in adjacent oilseed rape (Chapter VI) and winter cereal fields (Chapter VII). I recorded biodiversity and pest control potentials using a variety of different methods (Chapters II, V, VI & VII). Moreover, I validated the methodology I used to assess predator assemblages and predation rates (Chapters III & IV).
Chapter III: How to record ground dwelling predators?
Testing methodology is critical as it ensures scientific standards and trustworthy results. Pitfall traps are widely used to record ground dwelling predators, but little is known about how different trap types affect catches. I compared different types of pitfall traps that had been used in previous studies in respect to resulting carabid beetle assemblages. While barrier traps collected more species and deliver more complete species inventories, conventional simple pitfall traps provide reliable results with comparatively little handling effort. Placing several simple pitfall traps in the field can compensate the difference while still saving handling effort.
Chapter IV: How to record predation rates?
A plethora of methods has been proposed and used for recording predation rates, but these have rarely been validated before use. I assessed whether a novel approach to record predation, the use of sentinel prey cards with glued on aphids, delivers realistic results. I compared different sampling efforts and showed that obtained predation rates were similar and could be linked to predator (carabid beetle) densities and body-sizes (a proxy often used for food intake rates). Thus, the method delivers reliable and meaningful predation rates.
Chapter V: Do AES habitats benefit multi-taxa biodiversity?
The main goal of AES is the conservation of biodiversity in agricultural landscapes. I investigated how effectively AES habitats with different temporal continuity fulfil this goal in differently structured landscapes. The different AES habitats investigated had variable effects on local biodiversity. Temporal continuity of AES habitats was the most important predictor with older, more temporally continuous habitats harbouring higher overall biodiversity and different species assemblages in most taxonomic groups than younger AES habitats. Results however varied among taxonomic groups and natural enemies were equally supported by younger habitats. Semi-natural habitats in the surrounding landscape and AES habitat size were of minor importance for local biodiversity and had limited effects. This stresses that newly established AES habitats alone cannot restore farmland biodiversity. Both AES habitats as well as more continuous semi-natural habitats synergistically increase overall biodiversity in agricultural landscapes.
Chapter VI: The effects of AES habitats on predators in adjacent oilseed rape fields
Apart from biodiversity conservation, ensuring ecosystem service delivery in agricultural landscapes is a crucial goal of AES. I therefore investigated the effects of adjacent AES habitats on ground dwelling predator assemblages in oilseed rape fields. I found clear distance decay effects from the field edges into the field centres on both richness and densities of ground dwelling predators. Direct effects of adjacent AES habitats on assemblages in oilseed rape fields however were limited and only visible in functional traits of carabid beetle assemblages. Adjacent AES habitats doubled the proportion of predatory carabid beetles indicating a beneficial role for pest control. My results show that pest control potentials are largest close to the field edges and beneficial effects are comparably short ranged.
Chapter VII: The effects of AES habitats on pest control in adjacent cereal fields
Whether distance functions and potential effects of AES habitats are universal across crops is unknown. Therefore, I assessed distance functions of predators, pests, predation rates and yields after crop rotation in winter cereals using the same study design as in the previous year. Resulting distance functions were not uniform and differed from those found in oilseed rape in the previous year, indicating that the interactions between certain adjacent habitats vary with habitat and crop types. Distance functions of cereal-leaf beetles (important cereal pests) and parasitoid wasps were moreover modulated by semi-natural habitat proportion in the surrounding landscapes. Field edges buffered assemblage changes in carabid beetle assemblages over crop rotation confirming their important function as refuges for natural enemies. My results emphasize the beneficial role of field edges for pest control potentials. These findings back the calls for smaller field sizes and more diverse, more heterogeneously structured agricultural landscapes.
Chapter VIII: General Discussion
Countering biodiversity loss and ensuring ecosystem service provision in agricultural landscapes is intricate and requires strategic planning and restructuring of these landscapes. I showed that agricultural landscapes could benefit maximally from (i) a mixture of AES habitats and semi-natural habitats to support high levels of overall biodiversity and from (ii) smaller continuously managed agricultural areas (i.e. smaller field sizes or the insertion of AES elements within large fields) to maximize natural pest control potentials in crop fields. I propose a mosaic of younger AES habitats and semi-natural habitats to support ecosystem service providers and increase edge density for ecosystem service spillover into adjacent crops. The optimal extent and density of this network as well as the location in which AES and semi-natural habitats interact most beneficially with adjacent crops need further investigation. My results provide a further step towards more sustainable agricultural landscapes that simultaneously allow biodiversity to persist and maintain agricultural production under the framework of ecological intensification.
Since more than two centuries naturalists are fascinated by the profound changes in biodiversity observed along climatic gradients. Although the theories explaining changes in the diversity and the shape of organisms along climatic gradients belong to the foundations of modern ecology, our picture on the spatial patterns and drivers of biodiversity is far from being complete. Ambiguities in theory and data are common and past work has been strongly concentrated on plants and vertebrates. In the last two decades, interest in the fundamental processes structuring diversity along climatic gradients gained new impetus as they are expected to improve our understanding about how ecosystems will respond to global environmental changes. Global temperatures are rising faster than ever before; natural habitats are transformed into agricultural land and existing land use systems get more and more intensified to meet the demands of growing human populations. The fundamental shifts in the abiotic and biotic environment are proclaimed to affect ecosystems all over the world; however, precise predictions about how ecosystems respond to global changes are still lacking. We investigated diversity, traits and ecosystem services of wild bees along climate and land use gradients on Mount Kilimanjaro (Tanzania, East Africa). Wild bees play a major role in ecosystems, as they contribute to the reproduction and performance of wild and crop plants. Their responsiveness to environmental changes is therefore of high ecological and economic importance.
Temperature and energy resources have often been suggested to be the main determinants of global and local species richness, but the mechanisms behind remain poorly understood. In the study described in chapter II we analyzed species richness patterns of wild bees along climate and land use gradients on Mount Kilimanjaro and disentangled the factors explaining most of the changes in bee richness. We found that floral resources had a weak but significant effect on pollinator abundance, which in turn was positively related to species richness. However, temperature was the strongest predictor of species richness, affecting species richness both directly and indirectly by positively influencing bee abundances. We observed higher levels of bee-flower-interactions at higher temperatures, independently of flower and bee abundances. This suggests that temperature restricts species richness by constraining the exploitation of resources by ectotherms. Current land use did not negatively affect species richness. We conclude that the richness of bees is explained by both temperature and resource availability, whereas temperature plays the dominant role as it limits the access of ectotherms to floral resources and may accelerate ecological and evolutionary processes that drive the maintenance and origination of diversity.
Not only species numbers, but also morphological traits like body size are expected to be shaped by both physiological and energetic constraints along elevational gradients. Paradoxically, Bergmann´s rule predicts increases of body sizes in cooler climates resulting from physiological constraints, while species-energy theory suggests declines in the mean body size of species caused by increased extinction probabilities for large-bodied species in low-energy habitats. In chapter III we confronted this ambiguity with field data by studying community-wide body size variation of wild bees on Mt. Kilimanjaro. We found that along a 3680 m elevational gradient bee individuals became on average larger within species, while large species were increasingly absent from high-elevational communities. This demonstrates, on the one hand, how well-established, but apparently contrasting ecological theories can be merged through the parallel consideration of different levels of biological organization. On the other hand it signals that the extinction risk in the course of environmental change is not equally distributed among species within a community.
Land use intensification is known to threaten biodiversity, but the consequences for ecosystem services are still a matter of debate. In chapter IV, we experimentally tested the single and combined contributions of pest predators and pollinators to coffee production along a land use intensification gradient on Mount Kilimanjaro. We found that pest predation increased fruit set by on average 9%, while pollination increased fruit weight of coffee by on average 7.4%. Land use had no significant effect on both ecosystem services. However, we found that in coffee plantations with most intensified land use, pollination services were virtually exclusively provided by the honey bee (Apis mellifera). The reliance on a single pollinator species is risky, as possible declines of that species may directly lower pollination services, resulting in yield losses. In contrast, pollination services in structurally complex homegardens were found to be provided by a diverse pollinator community, increasing the stability of pollination services in a long term.
We showed that on Mount Kilimanjaro pollinator communities changed along elevational gradients in terms of species richness (chapter II) and trait composition (chapter III). Temperature and the temperature-mediated accessibility of resources were identified as important predictors of these patterns, which contributes to our fundamental understanding about the factors that shape ectothermic insect communities along climatic gradients. The strong temperature-dependence of pollinators suggests that temperature shifts in the course of global change are likely to affect pollinator communities. Pollinators might either profit from rising temperatures, or shift to higher elevations, which could result in related biotic attrition in the lowland with consequences for the provision of ecosystem services in cropping systems. Up to now, land use intensification had no significant impact on the diversity of pollinator communities and their ecosystem services. Pollinators might profit from the strong landscape heterogeneity in the region and from the amount of flower resources in the understory of cropping systems. However,progressing homogenization of the landscape and the pronounced application of pesticides could result in reduced diversity and dominance of single species, as we already found in sun coffee plantations. Such shifts in community compositions could threaten the stability of ecosystem services within cropping and natural systems in a long term.
1. Today honey bee colonies face a wide range of challenges in modern agricultural landscapes which entails the need for a comprehensive investigation of honey bees in a landscape context and the assessment of environmental risks. Within this dissertation the pollen foraging of honey bee colonies is studied in different agricultural landscapes to gain insight into the use of pollen resources and the influence of landscape structure across the season. General suggestions for landscape management to support honey bees and other pollinators are derived.
2. Decoding of waggle dances and a subsequent spatial foraging analysis are used as methods in Chapters 4 and 5 to study honey bee colonies in agricultural landscapes. The recently developed metabarcoding of mixed pollen samples was applied for the first time in honey bee foraging ecology and allowed for a detailed analysis of pollen, that was trapped from honey bees in front hive entrances (Chapter 6).
3. Pollen identification through molecular sequencing and DNA barcoding has been proposed as an alternative approach to light microscopy, which still is a tedious and error-prone task. In this study we assessed mixed pollen probes through next-generation sequencing and developed a bioinformatic workflow to analyse these high-throughput data with a newly created reference database. To evaluate the feasibility, we compared results from classical identification based on light microscopy from the same samples with our sequencing results. Abundance estimations from sequencing data were significantly correlated with counted abundances through light microscopy. Next-generation sequencing thus presents a useful and efficient workflow to identify pollen at the genus and species level without requiring specialized palynological expert knowledge.
4. During maize flowering, four observation hives were placed in and rotated between 11 landscapes covering a gradient in maize acreage. A higher foraging frequency on maize fields compared to other landuse types showed that maize is an intensively used pollen resource for honey bee colonies. Mean foraging distances were significantly shorter for maize pollen than for other pollen origins, indicating that effort is put into collecting a diverse pollen diet. The percentage of maize pollen foragers did not increase with maize acreage in the landscape and was not reduced by grassland area as an alternative pollen resource. Our findings allow estimating the distance-related exposure risk of honey bee colonies to pollen from surrounding maize fields treated with systemic insecticides.
5. It is unknown how an increasing area of mass-flowering crops like oilseed rape (OSR) or a decrease of semi-natural habitats (SNH) change the temporal and spatial availability of pollen resources for honey bee colonies, and thus foraging distances and frequency in different habitat types. Sixteen observation hives were placed in and rotated between 16 agricultural landscapes with independent gradients of OSR and SNH area within 2 km to analyze foraging distances and frequencies. SNH and OSR reduced foraging distance at different spatial scales and depending on season, with possible benefits for the performance of honey bee colonies. Frequency of pollen foragers per habitat type was equally high for SNH, grassland and OSR fields, but lower for other crops and forest. In landscapes with a small proportion of SNH a significantly higher density of pollen foragers on SNH was observed, indicating the limitation of pollen resources in simple agricultural landscapes and the importance of SNH.
6. Quantity and diversity of collected pollen can influence the growth and health of honey bee colonies, but little is known about the influence of landscape structure on pollen diet. In a field experiment we rotated 16 honey bee colonies across 16 agricultural landscapes (see also Chapter 5), used traps to get samples of collected pollen and observed the intra-colonial dance communication to gain information about foraging distances. Neither the amount of collected pollen nor pollen diversity were related to landscape diversity. The revealed increase of foraging distances with decreasing landscape diversity suggests that honey bees compensate for a lower landscape diversity by increasing their pollen foraging range in order to maintain pollen amount and diversity.
7. Our results show the importance of diverse pollen resources for honey bee colonies in agricultural landscapes. Beside the risk of exposure to pesticides honey bees face the risk of nutritional deficiency with implications for their health. By modifying landscape composition and therefore availability of resources we are able to contribute to the wellbeing of honey bees. Agri-environmental schemes aiming to support pollinators should focus on possible spatial and temporal gaps in pollen availability and diversity in agricultural landscapes.
Accurate crop monitoring in response to climate change at a regional or field scale
plays a significant role in developing agricultural policies, improving food security,
forecasting, and analysing global trade trends. Climate change is expected to
significantly impact agriculture, with shifts in temperature, precipitation patterns, and
extreme weather events negatively affecting crop yields, soil fertility, water availability,
biodiversity, and crop growing conditions. Remote sensing (RS) can provide valuable
information combined with crop growth models (CGMs) for yield assessment by
monitoring crop development, detecting crop changes, and assessing the impact of
climate change on crop yields. This dissertation aims to investigate the potential of RS
data on modelling long-term crop yields of winter wheat (WW) and oil seed rape (OSR)
for the Free State of Bavaria (70,550 km2
), Germany. The first chapter of the dissertation
describes the reasons favouring the importance of accurate crop yield predictions for
achieving sustainability in agriculture. Chapter second explores the accuracy
assessment of the synthetic RS data by fusing NDVIs of two high spatial resolution data
(high pair) (Landsat (30 m, 16-days; L) and Sentinel-2 (10 m, 5–6 days; S), with four low
spatial resolution data (low pair) (MOD13Q1 (250 m, 16-days), MCD43A4 (500 m, one
day), MOD09GQ (250 m, one-day), and MOD09Q1 (250 m, 8-days)) using the spatial
and temporal adaptive reflectance fusion model (STARFM), which fills regions' cloud
or shadow gaps without losing spatial information. The chapter finds that both L-MOD13Q1 (R2 = 0.62, RMSE = 0.11) and S-MOD13Q1 (R2 = 0.68, RMSE = 0.13) are more
suitable for agricultural monitoring than the other synthetic products fused. Chapter
third explores the ability of the synthetic spatiotemporal datasets (obtained in chapter
2) to accurately map and monitor crop yields of WW and OSR at a regional scale. The
chapter investigates and discusses the optimal spatial (10 m, 30 m, or 250 m), temporal
(8 or 16-day) and CGMs (World Food Studies (WOFOST), and the semi-empiric light
use efficiency approach (LUE)) for accurate crop yield estimations of both crop types.
Chapter third observes that the observations of high temporal resolution (8-day)
products of both S-MOD13Q1 and L-MOD13Q1 play a significant role in accurately
measuring the yield of WW and OSR. The chapter investigates that the simple light use
efficiency (LUE) model (R2 = 0.77 and relative RMSE (RRMSE) = 8.17%) that required fewer input parameters to simulate crop yield is highly accurate, reliable, and more
precise than the complex WOFOST model (R2 = 0.66 and RRMSE = 11.35%) with higher
input parameters. Chapter four researches the relationship of spatiotemporal fusion
modelling using STRAFM on crop yield prediction for WW and OSR using the LUE
model for Bavaria from 2001 to 2019. The chapter states the high positive correlation
coefficient (R) = 0.81 and R = 0.77 between the yearly R2 of synthetic accuracy and
modelled yield accuracy for WW and OSR from 2001 to 2019, respectively. The chapter
analyses the impact of climate variables on crop yield predictions by observing an
increase in R2
(0.79 (WW)/0.86 (OSR)) and a decrease in RMSE (4.51/2.57 dt/ha) when
the climate effect is included in the model. The fifth chapter suggests that the coupling
of the LUE model to the random forest (RF) model can further reduce the relative root
mean square error (RRMSE) from -8% (WW) and -1.6% (OSR) and increase the R2 by
14.3% (for both WW and OSR), compared to results just relying on LUE. The same
chapter concludes that satellite-based crop biomass, solar radiation, and temperature
are the most influential variables in the yield prediction of both crop types. Chapter six
attempts to discuss both pros and cons of RS technology while analysing the impact of
land use diversity on crop-modelled biomass of WW and OSR. The chapter finds that
the modelled biomass of both crops is positively impacted by land use diversity to the
radius of 450 (Shannon Diversity Index ~0.75) and 1050 m (~0.75), respectively. The
chapter also discusses the future implications by stating that including some dependent
factors (such as the management practices used, soil health, pest management, and
pollinators) could improve the relationship of RS-modelled crop yields with
biodiversity. Lastly, chapter seven discusses testing the scope of new sensors such as
unmanned aerial vehicles, hyperspectral sensors, or Sentinel-1 SAR in RS for achieving
accurate crop yield predictions for precision farming. In addition, the chapter highlights
the significance of artificial intelligence (AI) or deep learning (DL) in obtaining higher
crop yield accuracies.
Accurate crop monitoring in response to climate change at a regional or field scale plays a significant role in developing agricultural policies, improving food security, forecasting, and analysing global trade trends. Climate change is expected to significantly impact agriculture, with shifts in temperature, precipitation patterns, and extreme weather events negatively affecting crop yields, soil fertility, water availability, biodiversity, and crop growing conditions. Remote sensing (RS) can provide valuable information combined with crop growth models (CGMs) for yield assessment by monitoring crop development, detecting crop changes, and assessing the impact of climate change on crop yields. This dissertation aims to investigate the potential of RS data on modelling long-term crop yields of winter wheat (WW) and oil seed rape (OSR) for the Free State of Bavaria (70,550 km2), Germany. The first chapter of the dissertation describes the reasons favouring the importance of accurate crop yield predictions for achieving sustainability in agriculture. Chapter second explores the accuracy assessment of the synthetic RS data by fusing NDVIs of two high spatial resolution data (high pair) (Landsat (30 m, 16-days; L) and Sentinel-2 (10 m, 5–6 days; S), with four low spatial resolution data (low pair) (MOD13Q1 (250 m, 16-days), MCD43A4 (500 m, one day), MOD09GQ (250 m, one-day), and MOD09Q1 (250 m, 8-days)) using the spatial and temporal adaptive reflectance fusion model (STARFM), which fills regions' cloud or shadow gaps without losing spatial information. The chapter finds that both L-MOD13Q1 (R2 = 0.62, RMSE = 0.11) and S-MOD13Q1 (R2 = 0.68, RMSE = 0.13) are more suitable for agricultural monitoring than the other synthetic products fused. Chapter third explores the ability of the synthetic spatiotemporal datasets (obtained in chapter 2) to accurately map and monitor crop yields of WW and OSR at a regional scale. The chapter investigates and discusses the optimal spatial (10 m, 30 m, or 250 m), temporal (8 or 16-day) and CGMs (World Food Studies (WOFOST), and the semi-empiric light use efficiency approach (LUE)) for accurate crop yield estimations of both crop types. Chapter third observes that the observations of high temporal resolution (8-day) products of both S-MOD13Q1 and L-MOD13Q1 play a significant role in accurately measuring the yield of WW and OSR. The chapter investigates that the simple light use efficiency (LUE) model (R2 = 0.77 and relative RMSE (RRMSE) = 8.17%) that required fewer input parameters to simulate crop yield is highly accurate, reliable, and more precise than the complex WOFOST model (R2 = 0.66 and RRMSE = 11.35%) with higher input parameters. Chapter four researches the relationship of spatiotemporal fusion modelling using STRAFM on crop yield prediction for WW and OSR using the LUE model for Bavaria from 2001 to 2019. The chapter states the high positive correlation coefficient (R) = 0.81 and R = 0.77 between the yearly R2 of synthetic accuracy and modelled yield accuracy for WW and OSR from 2001 to 2019, respectively. The chapter analyses the impact of climate variables on crop yield predictions by observing an increase in R2 (0.79 (WW)/0.86 (OSR)) and a decrease in RMSE (4.51/2.57 dt/ha) when the climate effect is included in the model. The fifth chapter suggests that the coupling of the LUE model to the random forest (RF) model can further reduce the relative root mean square error (RRMSE) from -8% (WW) and -1.6% (OSR) and increase the R2 by 14.3% (for both WW and OSR), compared to results just relying on LUE. The same chapter concludes that satellite-based crop biomass, solar radiation, and temperature are the most influential variables in the yield prediction of both crop types. Chapter six attempts to discuss both pros and cons of RS technology while analysing the impact of land use diversity on crop-modelled biomass of WW and OSR. The chapter finds that the modelled biomass of both crops is positively impacted by land use diversity to the radius of 450 (Shannon Diversity Index ~0.75) and 1050 m (~0.75), respectively. The chapter also discusses the future implications by stating that including some dependent factors (such as the management practices used, soil health, pest management, and pollinators) could improve the relationship of RS-modelled crop yields with biodiversity. Lastly, chapter seven discusses testing the scope of new sensors such as unmanned aerial vehicles, hyperspectral sensors, or Sentinel-1 SAR in RS for achieving accurate crop yield predictions for precision farming. In addition, the chapter highlights the significance of artificial intelligence (AI) or deep learning (DL) in obtaining higher crop yield accuracies.
Anthropogenic activities are causing air pollution. Amongst air pollutants, tropospheric ozone is a major threat to human health and ecosystem functioning. In this dissertation, I present three studies that aimed at increasing our knowledge on how plant exposure to ozone affects its reproduction and its interactions with insect herbivores and pollinators.
For this purpose, a new fumigation system was built and placed in a greenhouse. The annual plant Sinapis arvensis (wild mustard) was used as the model plant.
Plants were exposed to either 0 ppb (control) or 120 ppb of ozone, for variable amounts of time and at different points of their life cycle. After fumigation, plants were exposed to herbivores or pollinators in the greenhouse, or to both groups of insects in the field.
My research shows that ozone affected reproductive performance differently, depending on the timing of exposure: plants exposed at earlier ages had their reproductive fitness increased, while plants exposed later in their life cycle showed a tendency for reduced reproductive fitness. Plant phenology was a key factor influencing reproductive fitness: ozone accelerated flowering and increased the number of flowers produced by plants exposed at early ages, while plants exposed to ozone at later ages tended to have fewer flowers. On the other hand, the ozone-mediated changes in plant-insect interactions had little impact on plant reproductive success.
The strongest effect of ozone on plant-pollinator interactions was the change in the number of flower visits received per plant, which was strongly linked to the number of open flowers. This means that, as a rule, exposure of plants to ozone early in the life cycle resulted in a higher number of pollinator visits, while exposure later in the life cycle resulted in fewer flower visits by potential pollinators. An exception was observed: the higher number of visits performed by large syrphid flies to young ozone-exposed plants than to the respective control plants went beyond the increase in the number of open flowers in those plants. Also, honeybees spent more time per flower in plants exposed to ozone than on control plants, while other pollinators spent similar amounts of time in control and ozone-exposed plants. This guild-dependent preference for ozone-exposed plants may be due to species-specific preferences related to changes in the quality and quantity of floral rewards.
In the field, ozone-exposed plants showed only a tendency for increased colonization by sucking herbivores and slightly more damage by chewing herbivores than control plants. On the other hand, in the greenhouse experiment, Pieris brassicae butterflies preferred control plants over ozone-exposed plants as oviposition sites. Eggs laid on ozone-exposed plants took longer to hatch, but the chances of survival were higher. Caterpillars performed better in control plants than in ozone-exposed plants, particularly when the temperature was high.
Most of the described effects were dependent on the duration and timing of the ozone exposure and the observed temperature, with the strongest effects being observed for longer exposures and higher temperatures. Furthermore, the timing of exposure altered the direction of the effects.
The expected climate change provides ideal conditions for further increases in tropospheric ozone concentrations, therefore for stronger effects on plants and plant-insect interactions. Acceleration of flowering caused by plant exposure to ozone may put plant-pollinator interactions at risk by promoting desynchronization between plant and pollinator activities. Reduced performance of caterpillars feeding on ozone-exposed plants may weaken herbivore populations. On the other hand, the increased plant reproduction that results from exposing young plants to ozone may be a source of good news in the field of horticulture, when similar results would be achieved in high-value crops. However, plant response to ozone is highly species-specific. In fact, Sinapis arvensis is considered a weed and the advantage conferred by ozone exposure may increase its competitiveness, with negative consequences for crops or plant communities in general. Overall, plant exposure to ozone might constitute a threat for the balance of natural and agro-ecosystems.
The importance of understanding species extinctions and its consequences for ecosystems and human life has been getting increasing public attention.
Nonetheless, regardless of how pressing the current biodiversity loss is, with rare exceptions, extinctions are actually not immediate.
Rather, they happen many generations after the disturbance that caused them.
This means that, at any point in time after a given disturbance, there is a number of extinctions that are expected to happen.
This number is the extinction debt.
As long as all the extinctions triggered by the disturbance have not happened, there is a debt to be paid.
This delay in extinctions can be interpreted as a window of opportunity, when conservation measures can be implemented.
In this thesis, I investigated the relative importance of ecological and evolutionary processes unfolding after different disturbances scenarios, to understand how this knowledge can be used to improve conservation practices aiming at controlling extinctions.
In the Introduction (chapter 1), I present the concept of extinction debts and the complicating factors behind its understanding.
Namely, I start by presenting i) the theoretical basis behind the definition of extinction debts, and how each theory informed different methodologies of study, ii) the complexity of understanding and predicting eco-evolutionary dynamics, and iii) the challenges to studying extinctions under a regime of widespread and varied disturbance of natural habitats.
I start the main body of the thesis (chapter 2) by summarizing the current state of empirical, theoretical, and methodological research on extinction debts.
In the last 10 years, extinction debts were detected all over the globe, for a variety of ecosystems and taxonomic groups.
When estimated - a rare occurrence, since quantifying debts requires often unavailable data - the sizes of these debts range from 9 to 90\% of current species richness and they have been sustained for periods ranging from 5 to 570 yr.
I identified two processes whose contributions to extinction debts have been studied more often, namely 1) life-history traits that prolong individual survival, and 2) population and metapopulation dynamics that maintain populations under deteriorated conditions.
Less studied are the microevolutionary dynamics happening during the payment of a debt, the delayed conjoint extinctions of interaction partners, and the extinction dynamics under different regimes of disturbances (e.g. habitat loss vs. climate change).
Based on these observations, I proposed a roadmap for future research to focus on these less studies aspects.
In chapters 3 and 4, I started to follow this roadmap.
In chapter 3, I used a genomically-explicit, individual-based model of a plant community to study the microevolutionary processes happening after habitat loss and climate change, and potentially contributing to the settlement of a debt.
I showed that population demographic recovery through trait adaptation, i.e. evolutionary rescue, is possible.
In these cases, rather than directional selection, trait change involved increase in trait variation, which I interpreted as a sign of disruptive selection.
Moreover, I disentangled evolutionary rescue from demographic rescue and show that the two types of rescue were equally important for community resistance, indicating that community re-assembly plays an important role in maintaining diversity following disturbance.
The results demonstrated the importance of accounting for eco-evolutionary processes at the community level to understand and predict biodiversity change.
Furthermore, they indicate that evolutionary rescue has a limited potential to avoid extinctions under scenarios of habitat loss and climate change.
In chapter 4, I analysed the effects of habitat loss and disruption of pollination function on the extinction dynamics of plant communities.
To do it, I used an individual, trait-based eco-evolutionary model (Extinction Dynamics Model, EDM) parameterized according to real-world species of calcareous grasslands.
Specifically, I compared the effects of these disturbances on the magnitude of extinction debts and species extinction times, as well as how species functional traits affect species survival.
I showed that the loss of habitat area generates higher number of immediate extinctions, but the loss of pollination generates higher extinction debt, as species take longer to go extinct.
Moreover, reproductive traits (clonal ability, absence of selfing and insect pollination) were the traits that most influenced the occurrence of species extinction as payment of the debt.
Thus, the disruption of pollination functions arose as a major factor in the creation of extinction debts.
Thus, restoration policies should aim at monitoring the status of this and other ecological processes and functions in undisturbed systems, to inform its re-establishment in disturbed areas.
Finally, I discuss the implications of these findings to i) the theoretical understanding of extinction debts, notably via the niche, coexistence, and metabolic theories, ii) the planning conservation measures, including communicating the very notion of extinction debts to improve understanding of the dimension of the current biodiversity crisis, and iii) future research, which must improve the understanding of the interplay between extinction cascades and extinction debts.
Chapter 1 – General introduction
Anthropogenic land-use and climate change are the major drivers of the global biodiversity loss. Yet, biodiversity is essential for human well-being, as we depend on the availability of potable water, sufficient food and further benefits obtained from nature. Each species makes a somewhat unique contribution to these ecosystem services. Furthermore, species tolerate environmental stressors, such as climate change, differently. Thus, biodiversity is both the "engine" and the "insurance" for human well-being in a changing climate. Here, I investigate the effects of temperature and land use on herbivory (Chapter 2), predation (Chapter 3) and pest control (Chapter 4), and at the same time identify features of habitats (e.g. plant richness, proximity to different habitat types) and landscapes (e.g. landscape diversity, proportion of oilseed rape area) as potential management targets in an adaptation strategy to climate change. Finally, I discuss the similarities and differences between factors influencing herbivory, predation and pest control, while placing the observations in the context of climate change as a multifaceted phenomenon, and highlighting starting points for sustainable insect pest management (Chapter 5).
Chapter 2 – Plant richness, land use and temperature differently shape invertebrate leaf-chewing herbivory on major plant functional groups
Invertebrate herbivores are temperature-sensitive. Rising temperatures increase their metabolic rates and thus their demand for carbon-rich relative to protein-rich resources, which can lead to changes in the diets of generalist herbivores. Here, we quantified leaf-area loss to chewing invertebrates among three plant functional groups (legumes, non-leguminous forbs and grasses), which largely differ in C:N (carbon:nitrogen) ratio. This reseach was conducted along spatial temperature and land-use gradients in open herbaceous vegetation adjacent to different habitat types (forest, grassland, arable field, settlement). Herbivory largely differed among plant functional groups and was higher on legumes than forbs and grasses, except in open areas in forests. There, herbivory was similar among plant functional groups and on legumes lower than in grasslands. Also the presence of many plant families lowered herbivory on legumes. This suggests that open areas in forests and diverse vegetation provide certain protection against leaf damage to some plant families (e.g. legumes). This could be used as part of a conservation strategy for protected species. Overall, the effects of the dominant habitat type in the vicinity and diverse vegetation outweighed those of temperature and large-scale land use (e.g. grassland proportion, landscape diversity) on herbivory of legumes, forbs and grasses at the present time.
Chapter 3 – Landscape diversity and local temperature, but not climate, affect arthropod predation among habitat types
Herbivorous insects underlie top-down regulation by arthropod predators. Thereby, predation rates depend on predator community composition and behaviour, which is shaped by temperature, plant richness and land use. How the interaction of these factors affects the regulatory performance of predators was unknown. Therefore, we assessed arthropod predation rates on artificial caterpillars along temperature, and land-use gradients. On plots with low local mean temperature (≤ 7°C) often not a single caterpillar was attacked, which may be due to the temperature-dependent inactivity of arthropods. However, multi-annual mean temperature, plant richness and the dominant habitat type in the vicinity did not substantially affect arthropod predation rates. Highest arthropod predation rates were observed in diverse landscapes (2-km scale) independently of the locally dominanting habitat type. As landscape diversity, but not multi-annual mean temperature, affected arthropod predation rates, the diversification of landscapes may also support top-down regulation of herbivores independent of moderate increases of multi-annual mean temperature in the near future.
Chapter 4 – Pest control and yield of winter oilseed rape depend on spatiotemporal crop-cover dynamics and flowering onset: implications for global warming
Winter oilseed rape is an important oilseed crop in Europe, yet its seed yield is diminished through pests such as the pollen beetle and stem weevils. Damage from pollen beetles depends on pest abundances, but also on the timing of infestation relative to crop development as the bud stage is particularly vulnerable. The development of both oilseed rape and pollen beetles is temperature-dependent, while temperature effects on pest abundances are yet unknown, which brings opportunities and dangers to oilseed rape cropping under increased temperatures. We obtained measures of winter oilseed rape (flowering time, seed yield) and two of its major pests (pollen beetle, stem weevils) for the first time along both land-use and temperature gradients. Infestation with stem weevils was not influenced by any temperature or land-use aspect considered, and natural pest regulation of pollen beetles in terms of parasitism rates of pollen beetle larvae was low (< 30%), except on three out of 29 plots. Nonetheless, we could identify conditions favouring low pollen beetle abundances per plant and high seed yields. Low pollen beetle densities were favoured by a constant oilseed rape area relative to the preceding year (5-km scale), whereas a strong reduction in area (> 40%) caused high pest densities (concentration effect). This occurred more frequently in warmer regions, due to drought around sowing, which contributed to increased pollen beetle numbers in those regions. Yet, in warmer regions, oilseed rape flowered early, which possibly led to partial escape from pollen beetle infestation in the most vulnerable bud stage. This is also suggested by higher seed yields of early flowering oilseed rape fields, but not per se at higher temperatures. Thus, early flowering (e.g. cultivar selection) and the interannual coordination of oilseed rape area offer opportunities for environmental-friendly pollen beetle management.
Chapter 5 – General discussion
Anthropogenic land-use and climate change are major threats to biodiversity, and consequently to ecosystem functions, although I could show that ecosystem functions such as herbivory and predation barely responded to temperature along a spatial gradient at present time. Yet, it is important to keep several points in mind: (i) The high rate of climate warming likely reduces the time that species will have to adapt to temperature in the future; (ii) Beyond mean temperatures, many aspects of climate will change; (iii) The compensation of biodiversity loss through functional redundancy in arthropod communities may be depleted at some point; (iv) Measures of ecosystem functions are limited by methodological filters, so that changes may be captured incompletely. Although much uncertainty of the effects of climate and land-use change on ecosystem functions remains, actions to halt biodiversity loss and to interfere with natural processes in an environmentally friendly way, e.g. reduction of herbivory on crops, are urgently needed. With this thesis, I contribute options to the environment-friendly regulation of herbivory, which are at least to some extent climate resilient, and at the same time make a contribution to halt biodiversity loss. Yet, more research and a transformation process is needed to make human action more sustainable. In terms of crop protection, this means that the most common method of treating pests with fast-acting pesticides is not necessarily the most sustainable. To realize sustainable strategies, collective efforts will be needed targeted at crop damage prevention through reducing pest populations and densities in the medium to long term. The sooner we transform human action from environmentally damaging to biodiversity promoting, the higher is our insurance asset that secures human well-being under a changing climate.
Despite belonging to the best described patterns in ecology, the mechanisms driving biodiversity along broad-scale climatic gradients, like the latitudinal gradient in diversity, remain poorly understood. Because of their high biodiversity, restricted spatial ranges, the continuous change in abiotic factors with altitude and their worldwide occurrence, mountains constitute ideal study systems to elucidate the predictors of global biodiversity patterns. However, mountain ecosystems are increasingly threatened by human land use and climate change. Since the consequences of such alterations on mountainous biodiversity and related ecosystem services are hardly known, research along elevational gradients is also of utmost importance from a conservation point of view. In addition to classical biodiversity research focusing on taxonomy, the significance of studying functional traits and their prominence in biodiversity ecosystem functioning (BEF) relationships is increasingly acknowledged. In this dissertation, I explore the patterns and drivers of mammal and dung beetle diversity along elevational and land use gradients on Mt. Kilimanjaro, Tanzania. Furthermore, I investigate the predictors of dung decomposition by dung beetles under different extinction scenarios.
Mammals are not only charismatic, they also fulfil important roles in ecosystems. They provide important ecosystem services such as seed dispersal and nutrient cycling by turning over high amounts of biomass. In chapter II, I show that mammal diversity and community biomass both exhibited a unimodal distribution with elevation on Mt.Kilimanjaro and were mainly impacted by primary productivity, a measure of the total food abundance, and the protection status of study plots. Due to their large size and endothermy, mammals, in contrast to most arthopods, are theoretically predicted to be limited by food availability. My results are in concordance with this prediction. The significantly higher diversity and biomass in the Kilimanjaro National Park and in other conservation areas underscore the important role of habitat protection is vital for the conservation of large mammal biodiversity on tropical mountains.
Dung beetles are dependent on mammals since they rely upon mammalian dung as a food and nesting resource. Dung beetles are also important ecosystem service providers: they play an important role in nutrient cycling, bioturbation, secondary seed dispersal and parasite suppression. In chapter III, I show that dung beetle diversity declined with elevation while dung beetle abundance followed a hump-shaped pattern along the elevational gradient. In contrast to mammals, dung beetle diversity was primarily predicted by temperature. Despite my attempt to accurately quantifiy mammalian dung resources by calculating mammalian defecation rates, I did not find an influence of dung resource availability on dung beetle richness. Instead, higher temperature translated into higher dung beetle diversity.
Apart from being important ecosystem service providers, dung beetles are also model organisms for BEF studies since they rely on a resource which can be quantified easily. In chapter IV, I explore dung decomposition by dung beetles along the elevational gradient by means of an exclosure experiment in the presence of the whole dung beetle community, in the absence of large dung beetles and without any dung beetles. I show that dung decomposition was the highest when the dung could be decomposed by the whole dung beetle community, while dung decomposition was significantly reduced in the sole presence of small dung beetles and the lowest in the absence of dung beetles. Furthermore, I demonstrate that the drivers of dung decomposition were depend on the intactness of the dung beetle community. While body size was the most important driver in the presence of the whole dung beetle community, species richness gained in importance when large dung beetles were excluded. In the most perturbed state of the system with no dung beetles present, temperature was the sole driver of dung decomposition. In conclusion, abiotic drivers become more important predictors of ecosystem services the more the study system is disturbed.
In this dissertation, I exemplify that the drivers of diversity along broad-scale climatic gradients on Mt. Kilimanjaro depend on the thermoregulatory strategy of organisms. While mammal diversity was mainly impacted by food/energy resources, dung beetle diversity was mainly limited by temperature. I also demonstrate the importance of protected areas for the preservation of large mammal biodiversity. Furthermore, I show that large dung beetles were disproportionately important for dung decomposition as dung decomposition significantly decreased when large dung beetles were excluded. As regards land use, I did not detect an overall effect on dung beetle and mammal diversity nor on dung beetle-mediated dung decomposition. However, for the most specialised mammal trophic guilds and dung beetle functional groups, negative land use effects were already visible. Even though the current moderate levels of land use on Mt. Kilimanjaro can sustain high levels of biodiversity, the pressure of the human population on Mt. Kilimanjaro is increasing and further land use intensification poses a great threat to biodiversity. In synergy wih land use, climate change is jeopardizing current patterns and levels of biodiversity with the potential to displace communities, which may have unpredictable consequences for ecosystem service provisioning in the future.
1. Pollination of sexually reproducing plants requires pollen transfer agents, which can be biotic, abiotic or a combination of biotic and abiotic agents. The dominance of one of pollination system in wild plant communities depends on climatic factors and/or degrees of anthropogenic influences, which have effects on pollinator diversity and pollination function. Anthropogenic activities and climate change are also considered as main causes of ongoing invasion of invasive species into wild and managed habitats which can bring up competition for pollinators with possible negative consequences for the reproduction of co-occurring native plant species.
2. The study aimed to determine pollination systems and pollination limitation of invasive and native plant communities in natural savannah between 870 – 1130 m and semi-natural (managed) grassland between 1300 – 1750 m above sea level; effects of flower density and pollinator abundance on seed production of cross-pollinated and self-pollinated plants; and relationships of bee abundance and the proportion of cross- pollinated plants at the southern slope of Mount Kilimanjaro, Tanzania.
3. Pollinator-exclusion, open pollination and supplemental hand-pollination treatments were applied to 27 plant species in savannah and grassland habitats. Flowers were counted in each clusters based upon their species. Pollinators were sampled by using pan traps. Information-theory-based multi-model averaging and generalized linear mixed effects models were used to identify and analyze the effects of flower density, pollinator abundance, pollination treatments and habitat types on seed production. Regression models were used to determine relationships of altitude with bee abundance, and with proportion of cross-pollinated plants.
4. My results show that mean seed numbers of native plants were significantly lower in pollinator-exclusion treatments than in open-pollination treatments, indicating their reliance on pollinators for reproductive success. In contrast, seed numbers of invasive plants were similar in pollinator-exclusion and open-pollination treatments, demonstrating an ability of reproduction without pollinators. Despite of higher levels of self-pollination in invasive plants, supplemental hand-pollination treatments revealed pollen limitation in grassland and marginally in savannah habitats. There were no significant difference in seed numbers between supplemental hand pollination and open pollination treatments of native plant communities in savannah and grassland, which indicates no pollination limitation in the studied ecological system for native communities. Besides, grassland plants produced comparatively more seeds than savannah plants, however seeds in grasslands were lighter than those of the savannah which may be due to nutrient limitation in grassland.
5. I found 12 cross-pollinated and 15 self-pollinated plants along altitudinal gradient after comparing seeds from pollinator-excluded and open-pollinated experiments. I also found that proportions of cross-pollinated plants and bee abundance simultaneously decreased with increasing altitude. All cross-pollinated plants were native and grew in savannah habitats, with an exception of one species.
6. Neither effects of focal flower density nor a significant interaction between focal flower densities and bee abundance for self-pollinated plants were observed. However, there were effects of focal flower densities and interactions of flower density with bee abundance for cross-pollinated plants. Non-focal flower density has no significant effects on seed production of cross-pollinated and self-pollinated plants.
7. The results show that native plants depend more on cross-pollination than invasive plants, despite of most native plants in managed habitat (grassland) rely on self-pollination for reproduction. The tendency of having more cross-pollinated plants in natural savannah which are in low altitude coincides with other finding that the cross-pollinated plants and bee abundance simultaneously decrease with increasing altitude. Therefore, our findings support the hypotheses that self-fertilization of flowering plants increases with increasing altitude, and pollinator limitation is most pronounced in managed or disturbed habitats. Despite of reduction of pollinators in grassland, only invasive plants experience pollen limitation, which may be due to poor integration with available pollinator networks.
8. I also found bee abundance and flower density are not the main pollination factors required by self-pollinated plants during reproduction. However, focal flower density, which influences pollinator diversity, is more applicable to cross-pollinated plants. Climate change and anthropogenic activities in natural habitats are factors that influence pollinator abundance and functioning, which lead to a shift of mating systems in plant communities so as to assure their reproduction.