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Animal movement is a crucial aspect of life, influencing ecological and evolutionary processes. It plays an important role in shaping biodiversity patterns, connecting habitats and ecosystems. Anthropogenic landscape changes, such as in agricultural environments, can impede the movement of animals by affecting their ability to locate resources during recurring movements within home ranges and, on a larger scale, disrupt migration or dispersal. Inevitably, these changes in movement behavior have far-reaching consequences on the mobile link functions provided by species inhabiting such extensively altered matrix areas. In this thesis, I investigate the movement characteristics and activity patterns of the European hare (Lepus europaeus), aiming to understand their significance as a pivotal species in fragmented agricultural landscapes. I reveal intriguing results that shed light on the importance of hares for seed dispersal, the influence of personality traits on behavior and space use, the sensitivity of hares to extreme weather conditions, and the impacts of GPS collaring on mammals' activity patterns and movement behavior.
In Chapter I, I conducted a controlled feeding experiment to investigate the potential impact of hares on seed dispersal. By additionally utilizing GPS data of hares in two contrasting landscapes, I demonstrated that hares play a vital role, acting as effective mobile linkers for many plant species in small and isolated habitat patches. The analysis of seed intake and germination success revealed that distinct seed traits, such as density, surface area, and shape, profoundly affect hares' ability to disperse seeds through endozoochory. These findings highlight the interplay between hares and plant communities and thus provide valuable insights into seed dispersal mechanisms in fragmented landscapes.
By employing standardized behavioral tests in Chapter II, I revealed consistent behavioral responses among captive hares while simultaneously examining the intricate connection between personality traits and spatial patterns within wild hare populations. This analysis provides insights into the ecological interactions and dynamics within hare populations in agricultural habitats. Examining the concept of animal personality, I established a link between personality traits and hare behavior. I showed that boldness, measured through standardized tests, influences individual exploration styles, with shy and bold hares exhibiting distinct space use patterns. In addition to providing valuable insights into the role of animal personality in heterogeneous environments, my research introduced a novel approach demonstrating the feasibility of remotely assessing personality types using animal-borne sensors without additional disturbance of the focal individual.
While climate conditions severely impact the activity and, consequently, the fitness of wildlife species across the globe, in Chapter III, I uncovered the sensitivity of hares to temperature, humidity, and wind speed during their peak reproduction period. I found a strong response in activity to high temperatures above 25°C, with a particularly pronounced effect during temperature extremes of over 35°C. The non-linear relationship between temperature and activity was characterized by contrasting responses observed for day and night. These findings emphasize the vulnerability of hares to climate change and the potential consequences for their fitness and population dynamics with the ongoing rise of temperature.
Since such insights can only be obtained through capturing and tagging free-ranging animals, I assessed potential impacts and the recovery process post-collar attachment in Chapter IV. For this purpose, I examined the daily distances moved and the temporal-associated activity of 1451 terrestrial mammals out of 42 species during their initial tracking period. The disturbance intensity and the speed of recovery varied across species, with herbivores, females, and individuals captured and collared in relatively secluded study areas experiencing more pronounced disturbances due to limited anthropogenic influences.
Mobile linkers are essential for maintaining biodiversity as they influence the dynamics and resilience of ecosystems. Furthermore, their ability to move through fragmented landscapes makes them a key component for restoring disturbed sites. Individual movement decisions determine the scale of mobile links, and understanding variations in space use among individuals is crucial for interpreting their functions. Climate change poses further challenges, with wildlife species expected to adjust their behavior, especially in response to high-temperature extremes, and comprehending the anthropogenic influence on animal movements will remain paramount to effective land use planning and the development of successful conservation strategies.
This thesis provides a comprehensive ecological understanding of hares in agricultural landscapes. My research findings underscore the importance of hares as mobile linkers, the influence of personality traits on behavior and spatial patterns, the vulnerability of hares to extreme weather conditions, and the immediate consequences of collar attachment on mammalian movements. Thus, I contribute valuable insights to wildlife conservation and management efforts, aiding in developing strategies to mitigate the impact of environmental changes on hare populations. Moreover, these findings enable the development of methodologies aimed at minimizing the impacts of collaring while also identifying potential biases in the data, thereby benefiting both animal welfare and the scientific integrity of localization studies.
Climate change of anthropogenic origin is affecting Earth’s biodiversity and therefore ecosystems and their services. High latitude ecosystems are even more impacted than the rest of Northern Hemisphere because of the amplified polar warming. Still, it is challenging to predict the dynamics of high latitude ecosystems because of complex interaction between abiotic and biotic components. As the past is the key to the future, the interpretation of past ecological changes to better understand ongoing processes is possible. In the Quaternary, the Pleistocene experienced several glacial and interglacial stages that affected past ecosystems. During the last Glacial, the Pleistocene steppe-tundra was covering most of unglaciated northern hemisphere and disappeared in parallel to the megafauna’s extinction at the transition to the Holocene (~11,700 years ago). The origin of the steppe-tundra decline is not well understood and knowledge on the mechanisms, which caused shifts in past communities and ecosystems, is of high priority as they are likely comparable to those affecting modern ecosystems. Lake or permafrost core sediments can be retrieved to investigate past biodiversity at transitions between glacial and interglacial stages. Siberia and Beringia were the origin of dispersal of the steppe-tundra, which make investigation this area of high priority. Until recently, macrofossils and pollen were the most common approaches. They are designed to reconstruct past composition changes but have limit and biases. Since the end of the 20th century, sedimentary ancient DNA (sedaDNA) can also be investigated. My main objectives were, by using sedaDNA approaches to provide scientific evidence of compositional and diversity changes in the Northern Hemisphere ecosystems at the transition between Quaternary glacial and interglacial stages.
In this thesis, I provide snapshots of entire ancient ecosystems and describe compositional changes between Quaternary glacial and interglacial stages, and confirm the vegetation composition and the spatial and temporal boundaries of the Pleistocene steppe-tundra. I identify a general loss of plant diversity with extinction events happening in parallel of megafauna’ extinction. I demonstrate how loss of biotic resilience led to the collapse of a previously well-established system and discuss my results in regards to the ongoing climate change. With further work to constrain biases and limits, sedaDNA can be used in parallel or even replace the more established macrofossils and pollen approaches as my results support the robustness and potential of sedaDNA to answer new palaeoecological questions such as plant diversity changes, loss and provide snapshots of entire ancient biota.
Extreme weather and climate events are one of the greatest dangers for present-day society. Therefore, it is important to provide reliable statements on what changes in extreme events can be expected along with future global climate change. However, the projected overall response to future climate change is generally a result of a complex interplay between individual physical mechanisms originated within the different climate subsystems. Hence, a profound understanding of these individual contributions is required in order to provide meaningful assessments of future changes in extreme events. One aspect of climate change is the recently observed phenomenon of Arctic Amplification and the related dramatic Arctic sea ice decline, which is expected to continue over the next decades. The question to what extent Arctic sea ice loss is able to affect atmospheric dynamics and extreme events over mid-latitudes has received a lot of attention over recent years and still remains a highly debated topic.
In this respect, the objective of this thesis is to contribute to a better understanding on the impact of future Arctic sea ice retreat on European temperature extremes and large-scale atmospheric dynamics.
The outcomes are based on model data from the atmospheric general circulation model ECHAM6. Two different sea ice sensitivity simulations from the Polar Amplification Intercomparison Project are employed and contrasted to a present day reference experiment: one experiment with prescribed future sea ice loss over the entire Arctic, as well as another one with sea ice reductions only locally prescribed over the Barents-Kara Sea.% prescribed over the entire Arctic, as well as only locally over the Barent/Karasea with a present day reference experiment.
The first part of the thesis focuses on how future Arctic sea ice reductions affect large-scale atmospheric dynamics over the Northern Hemisphere in terms of occurrence frequency changes of five preferred Euro-Atlantic circulation regimes. When compared to circulation regimes computed from ERA5 it shows that ECHAM6 is able to realistically simulate the regime structures. Both ECHAM6 sea ice sensitivity experiments exhibit similar regime frequency changes. Consistent with tendencies found in ERA5, a more frequent occurrence of a Scandinavian blocking pattern in midwinter is for instance detected under future sea ice conditions in the sensitivity experiments. Changes in occurrence frequencies of circulation regimes in summer season are however barely detected.
After identifying suitable regime storylines for the occurrence of European temperature extremes in winter, the previously detected regime frequency changes are used to quantify dynamically and thermodynamically driven contributions to sea ice-induced changes in European winter temperature extremes.
It is for instance shown how the preferred occurrence of a Scandinavian blocking regime under low sea ice conditions dynamically contributes to more frequent midwinter cold extreme occurrences over Central Europe. In addition, a reduced occurrence frequency of a Atlantic trough regime is linked to reduced winter warm extremes over Mid-Europe. Furthermore, it is demonstrated how the overall thermodynamical warming effect due to sea ice loss can result in less (more) frequent winter cold (warm) extremes, and consequently counteracts the dynamically induced changes.
Compared to winter season, circulation regimes in summer are less suitable as storylines for the occurrence of summer heat extremes.
Therefore, an approach based on circulation analogues is employed in order to quantify thermodyamically and dynamically driven contributions to sea ice-induced changes of summer heat extremes over three different European sectors. Reduced occurrences of blockings over Western Russia are detected in the ECHAM6 sea ice sensitivity experiments; however, arguing for dynamically and thermodynamically induced contributions to changes in summer heat extremes remains rather challenging.
While estimated numbers of past and future climate migrants are alarming, the growing empirical evidence suggests that the association between adverse climate-related events and migration is not universally positive. This dissertation seeks to advance our understanding of when and how climate migration emerges by analyzing heterogeneous climatic influences on migration in low- and middle-income countries. To this end, it draws on established economic theories of migration, datasets from physical and social sciences, causal inference techniques and approaches from systematic literature review. In three of its five chapters, I estimate causal effects of processes of climate change on inequality and migration in India and Sub-Saharan Africa. By employing interaction terms and by analyzing sub-samples of data, I explore how these relationships differ for various segments of the population. In the remaining two chapters, I present two systematic literature reviews. First, I undertake a comprehensive meta-regression analysis of the econometric climate migration literature to summarize general climate migration patterns and explain the conflicting findings. Second, motivated by the broad range of approaches in the field, I examine the literature from a methodological perspective to provide best practice guidelines for studying climate migration empirically. Overall, the evidence from this dissertation shows that climatic influences on human migration are highly heterogeneous. Whether adverse climate-related impacts materialize in migration depends on the socio-economic characteristics of the individual households, such as wealth, level of education, agricultural dependence or access to adaptation technologies and insurance. For instance, I show that while adverse climatic shocks are generally associated with an increase in migration in rural India, they reduce migration in the agricultural context of Sub-Saharan Africa, where the average wealth levels are much lower so that households largely cannot afford the upfront costs of moving. I find that unlike local climatic shocks which primarily enhance internal migration to cities and hence accelerate urbanization, shocks transmitted via agricultural producer prices increase migration to neighboring countries, likely due to the simultaneous decrease in real income in nearby urban areas. These findings advance our current understanding by showing when and how economic agents respond to climatic events, thus providing explicit contexts and mechanisms of climate change effects on migration in the future. The resulting collection of findings can guide policy interventions to avoid or mitigate any present and future welfare losses from climate change-related migration choices.
Global heat adaptation among urban populations and its evolution under different climate futures
(2022)
Heat and increasing ambient temperatures under climate change represent a serious threat to human health in cities. Heat exposure has been studied extensively at a global scale. Studies comparing a defined temperature threshold with the future daytime temperature during a certain period of time, had concluded an increase in threat to human health. Such findings however do not explicitly account for possible changes in future human heat adaptation and might even overestimate heat exposure. Thus, heat adaptation and its development is still unclear. Human heat adaptation refers to the local temperature to which populations are adjusted to. It can be inferred from the lowest point of the U- or V-shaped heat-mortality relationship (HMR), the Minimum Mortality Temperature (MMT). While epidemiological studies inform on the MMT at the city scale for case studies, a general model applicable at the global scale to infer on temporal change in MMTs had not yet been realised. The conventional approach depends on data availability, their robustness, and on the access to daily mortality records at the city scale. Thorough analysis however must account for future changes in the MMT as heat adaptation happens partially passively. Human heat adaptation consists of two aspects: (1) the intensity of the heat hazard that is still tolerated by human populations, meaning the heat burden they can bear and (2) the wealth-induced technological, social and behavioural measures that can be employed to avoid heat exposure. The objective of this thesis is to investigate and quantify human heat adaptation among urban populations at a global scale under the current climate and to project future adaptation under climate change until the end of the century. To date, this has not yet been accomplished. The evaluation of global heat adaptation among urban populations and its evolution under climate change comprises three levels of analysis. First, using the example of Germany, the MMT is calculated at the city level by applying the conventional method. Second, this thesis compiles a data pool of 400 urban MMTs to develop and train a new model capable of estimating MMTs on the basis of physical and socio-economic city characteristics using multivariate non-linear multivariate regression. The MMT is successfully described as a function of the current climate, the topography and the socio-economic standard, independently of daily mortality data for cities around the world. The city-specific MMT estimates represents a measure of human heat adaptation among the urban population. In a final third analysis, the model to derive human heat adaptation was adjusted to be driven by projected climate and socio-economic variables for the future. This allowed for estimation of the MMT and its change for 3 820 cities worldwide for different combinations of climate trajectories and socio-economic pathways until 2100. The knowledge on the evolution of heat adaptation in the future is a novelty as mostly heat exposure and its future development had been researched. In this work, changes in heat adaptation and exposure were analysed jointly. A wide range of possible health-related outcomes up to 2100 was the result, of which two scenarios with the highest socio-economic developments but opposing strong warming levels were highlighted for comparison. Strong economic growth based upon fossil fuel exploitation is associated with a high gain in heat adaptation, but may not be able to compensate for the associated negative health effects due to increased heat exposure in 30% to 40% of the cities investigated caused by severe climate change. A slightly less strong, but sustainable growth brings moderate gains in heat adaptation but a lower heat exposure and exposure reductions in 80% to 84% of the cities in terms of frequency (number of days exceeding the MMT) and intensity (magnitude of the MMT exceedance) due to a milder global warming. Choosing a 2 ° C compatible development by 2100 would therefore lower the risk of heat-related mortality at the end of the century. In summary, this thesis makes diverse and multidisciplinary contributions to a deeper understanding of human adaptation to heat under the current and the future climate. It is one of the first studies to carry out a systematic and statistical analysis of urban characteristics which are useful as MMT drivers to establish a generalised model of human heat adaptation, applicable at the global level. A broad range of possible heat-related health options for various future scenarios was shown for the first time. This work is of relevance for the assessment of heat-health impacts in regions where mortality data are not accessible or missing. The results are useful for health care planning at the meso- and macro-level and to urban- and climate change adaptation planning. Lastly, beyond having met the posed objective, this thesis advances research towards a global future impact assessment of heat on human health by providing an alternative method of MMT estimation, that is spatially and temporally flexible in its application.
The ongoing climate change is altering the living conditions for many organisms on this planet at an unprecedented pace. Hence, it is crucial for the survival of species to adapt to these changing conditions. In this dissertation Silene vulgaris is used as a model organism to understand the adaption strategies of widely distributed plant species to the current climate change. Especially plant species that possess a wide geographic range are expected to have a high phenotypic plasticity or to show genetic differentiation in response to the different climate conditions they grow in. However, they are often underrepresented in research.
In the greenhouse experiment presented in this thesis, I examined the phenotypic responses and plasticity in S. vulgaris to estimate its’ adaptation potential. Seeds from 25 wild European populations were collected along a latitudinal gradient and grown in a greenhouse under three different precipitation (65 mm, 75 mm, 90 mm) and two different temperature regimes (18°C, 21°C) that resembled a possible climate change scenario for central Europe. Afterwards different biomass and fecundity-related plant traits were measured.
The treatments significantly influenced the plants but did not reveal a latitudinal difference in response to climate treatments for most plant traits. The number of flowers per individual however, showed a stronger plasticity in northern European populations (e.g., Swedish populations) where numbers decreased more drastically with increased temperature and decreased precipitation.
To gain an even deeper understanding of the adaptation of S. vulgaris to climate change it is also important to reveal the underlying phylogeny of the sampled populations. Therefore, I analysed their population genetic structure through whole genome sequencing via ddRAD.
The sequencing revealed three major genetic clusters in the S. vulgaris populations sampled in Europe: one cluster comprised Southern European populations, one cluster Western European populations and another cluster contained central European populations. A following analysis of experimental trait responses among the clusters to the climate-change scenario showed that the genetic clusters significantly differed in biomass-related traits and in the days to flowering. However, half of the traits showed parallel response patterns to the experimental climate-change scenario.
In addition to the potential geographic and genetic adaptation differences to climate change this dissertation also deals with the response differences between the sexes in S. vulgaris. As a gynodioecious species populations of S. vulgaris consist of female and hermaphrodite
individuals and the sexes can differ in their morphological traits which is known as sexual dimorphism. As climate change is becoming an important factor influencing plant morphology it remains unclear if and how different sexes may respond in sexually dimorphic species. To examine this question the sex of each individual plant was determined during the greenhouse experiment and the measured plant traits were analysed accordingly. In general, hermaphrodites had a higher number of flowers but a lower number of leaves than females. With regards to the climate change treatment, I found that hermaphrodites showed a milder negative response to higher temperatures in the number of flowers produced and in specific leaf area (SLA) compared to females.
Synthesis – The significant treatment response in Silene vulgaris, independent of population origin in most traits suggests a high degree of universal phenotypic plasticity. Also, the three European intraspecific genetic lineages detected showed comparable parallel response patterns in half of the traits suggesting considerable phenotypic plasticity. Hence, plasticity might represent a possible adaptation strategy of this widely distributed species during ongoing and future climatic changes. The results on sexual dimorphism show that females and hermaphrodites are differing mainly in their number of flowers and females are affected more strongly by the experimental climate-change scenario. These results provide a solid knowledge basis on the sexual dimorphism in S. vulgaris under climate change, but further research is needed to determine the long-term impact on the breeding system for the species.
In summary this dissertation provides a comprehensive insight into the adaptation mechanisms and consequences of a widely distributed and gynodioecious plant species and leverages our understanding of the impact of anthropogenic climate change on plants.
Weather extremes pose a persistent threat to society on multiple layers. Besides an average of ~37,000 deaths per year, climate-related disasters cause destroyed properties and impaired economic activities, eroding people's livelihoods and prosperity. While global temperature rises – caused by anthropogenic greenhouse gas emissions – the direct impacts of climatic extreme events increase and will further intensify without proper adaptation measures. Additionally, weather extremes do not only have local direct effects. Resulting economic repercussions can propagate either upstream or downstream along trade chains causing indirect effects. One approach to analyze these indirect effects within the complex global supply network is the agent-based model Acclimate. Using and extending this loss-propagation model, I focus in this thesis on three aspects of the relation between weather extremes and economic repercussions.
First, extreme weather events cause direct impacts on local economic performance. I compute daily local direct output loss time series of heat stress, river floods, tropical cyclones, and their consecutive occurrence using (near-future) climate projection ensembles. These regional impacts are estimated based on physical drivers and local productivity distribution. Direct effects of the aforementioned disaster categories are widely heterogeneous concerning regional and temporal distribution. As well, their intensity changes differently under future warming. Focusing on the hurricane-impacted capital, I find that long-term growth losses increase with higher heterogeneity of a shock ensemble.
Second, repercussions are sectorally and regionally distributed via economic ripples within the trading network, causing higher-order effects. I use Acclimate to identify three phases of those economic ripples. Furthermore, I compute indirect impacts and analyze overall regional and global production and consumption changes. Regarding heat stress, global consumer losses double while direct output losses increase by a factor 1.5 between 2000 – 2039. In my research I identify the effect of economic ripple resonance and introduce it to climate impact research. This effect occurs if economic ripples of consecutive disasters overlap, which increases economic responses such as an enhancement of consumption losses. These loss enhancements can even be more amplified with increasing direct output losses, e.g. caused by climate crises.
Transport disruptions can cause economic repercussions as well. For this, I extend the model Acclimate with a geographical transportation route and expand the decision horizon of economic agents. Using this, I show that policy-induced sudden trade restrictions (e.g. a no-deal Brexit) can significantly reduce the longer-term economic prosperity of affected regions. Analyses of transportation disruptions in typhoon seasons indicate that severely affected regions must reduce production as demand falls during a storm. Substituting suppliers may compensate for fluctuations at the beginning of the storm, which fails for prolonged disruptions.
Third, possible coping mechanisms and adaptation strategies arise from direct and indirect economic responses to weather extremes. Analyzing annual trade changes due to typhoon-induced transport disruptions depict that overall exports rise. This trade resilience increases with higher network node diversification. Further, my research shows that a basic insurance scheme may diminish hurricane-induced long-term growth losses due to faster reconstruction in disasters aftermaths. I find that insurance coverage could be an economically reasonable coping scheme towards higher losses caused by the climate crisis. Indirect effects within the global economic network from weather extremes indicate further adaptation possibilities. For one, diversifying linkages reduce the hazard of sharp price increases. Next to this, close economic interconnections with regions that do not share the same extreme weather season can be economically beneficial in the medium run. Furthermore, economic ripple resonance effects should be considered while computing costs. Overall, an increase in local adaptation measures reduces economic ripples within the trade network and possible losses elsewhere. In conclusion, adaptation measures are necessary and potential present, but it seems rather not possible to avoid all direct or indirect losses.
As I show in this thesis, dynamical modeling gives valuable insights into how direct and indirect economic impacts arise from different categories of weather extremes. Further, it highlights the importance of resolving individual extremes and reflecting amplifying effects caused by incomplete recovery or consecutive disasters.
River floods are among the most devastating natural hazards worldwide. As their generation is highly dependent on climatic conditions, their magnitude and frequency are projected to be affected by future climate change. Therefore, it is crucial to study the ways in which a changing climate will, and already has, influenced flood generation, and thereby flood hazard. Additionally, it is important to understand how other human influences - specifically altered land cover - affect flood hazard at the catchment scale.
The ways in which flood generation is influenced by climatic and land cover conditions differ substantially in different regions. The spatial variability of these effects needs to be taken into account by using consistent datasets across large scales as well as applying methods that can reflect this heterogeneity. Therefore, in the first study of this cumulative thesis a complex network approach is used to find 10 clusters of similar flood behavior among 4390 catchments in the conterminous United States. By using a consistent set of 31 hydro-climatological and land cover variables, and training a separate Random Forest model for each of the clusters, the regional controls on flood magnitude trends between 1960-2010 are detected. It is shown that changes in rainfall are the most important drivers of these trends, while they are regionally controlled by land cover conditions.
While climate change is most commonly associated with flood magnitude trends, it has been shown to also influence flood timing. This can lead to trends in the size of the area across which floods occur simultaneously, the flood synchrony scale. The second study is an analysis of data from 3872 European streamflow gauges and shows that flood synchrony scales have increased in Western Europe and decreased in Eastern Europe. These changes are attributed to changes in flood generation, especially a decreasing relevance of snowmelt. Additionally, the analysis shows that both the absolute values and the trends of flood magnitudes and flood synchrony scales are positively correlated. If these trends persist in the future and are not accounted for, the combined increases of flood magnitudes and flood synchrony scales can exceed the capacities of disaster relief organizations and insurers.
Hazard cascades are an additional way through which climate change can influence different aspects of flood hazard. The 2019/2020 wildfires in Australia, which were preceded by an unprecedented drought and extinguished by extreme rainfall that led to local flooding, present an opportunity to study the effects of multiple preceding hazards on flood hazard. All these hazards are individually affected by climate change, additionally complicating the interactions within the cascade. By estimating and analyzing the burn severity, rainfall magnitude, soil erosion and stream turbidity in differently affected tributaries of the Manning River catchment, the third study shows that even low magnitude floods can pose a substantial hazard within a cascade.
This thesis shows that humanity is affecting flood hazard in multiple ways with spatially and temporarily varying consequences, many of which were previously neglected (e.g. flood synchrony scale, hazard cascades). To allow for informed decision making in risk management and climate change adaptation, it will be crucial to study these aspects across the globe and to project their trajectories into the future. The presented methods can depict the complex interactions of different flood drivers and their spatial variability, providing a basis for the assessment of future flood hazard changes. The role of land cover should be considered more in future flood risk modelling and management studies, while holistic, transferable frameworks for hazard cascade assessment will need to be designed.
The evolution of life on Earth has been driven by disturbances of different types and magnitudes over the 4.6 million years of Earth’s history (Raup, 1994, Alroy, 2008). One example for such disturbances are mass extinctions which are characterized by an exceptional increase in the extinction rate affecting a great number of taxa in a short interval of geologic time (Sepkoski, 1986). During the 541 million years of the Phanerozoic, life on Earth suffered five exceptionally severe mass extinctions named the “Big Five Extinctions”. Many mass extinctions are linked to changes in climate
(Feulner, 2009). Hence, the study of past mass extinctions is not only intriguing, but can also provide insights into the complex nature of the Earth system. This thesis aims at deepening our understanding of the triggers of mass extinctions and how they affected life. To accomplish this, I investigate changes in climate during two of the Big Five extinctions using a coupled climate model.
During the Devonian (419.2–358.9 million years ago) the first vascular plants and vertebrates evolved on land while extinction events occurred in the ocean (Algeo et al., 1995). The causes of these formative changes, their interactions and their links to changes in climate are still poorly understood. Therefore, we explore the sensitivity of the Devonian climate to various boundary conditions using an intermediate-complexity climate model (Brugger et al., 2019). In contrast to Le Hir et al. (2011), we find only a minor biogeophysical effect of changes in vegetation cover due to unrealistically high soil albedo values used in the earlier study. In addition, our results cannot support the strong influence of orbital parameters on the Devonian climate, as simulated with a climate model with a strongly simplified ocean model (De Vleeschouwer et al., 2013, 2014, 2017). We can only reproduce the changes in Devonian climate suggested by proxy data by decreasing atmospheric CO2. Still, finding agreement between the evolution of sea surface temperatures reconstructed from proxy data (Joachimski et al., 2009) and our simulations remains challenging and suggests a lower δ18O ratio of Devonian seawater. Furthermore, our study of the sensitivity of the Devonian climate reveals a prevailing mode of climate variability on a timescale of decades to centuries. The quasi-periodic ocean temperature fluctuations are linked to a physical mechanism of changing sea-ice cover, ocean convection and overturning in high northern latitudes.
In the second study of this thesis (Dahl et al., under review) a new reconstruction of atmospheric CO2 for the Devonian, which is based on CO2-sensitive carbon isotope fractionation in the earliest vascular plant fossils, suggests a much earlier drop of atmo- spheric CO2 concentration than previously reconstructed, followed by nearly constant CO2 concentrations during the Middle and Late Devonian. Our simulations for the Early Devonian with identical boundary conditions as in our Devonian sensitivity study (Brugger et al., 2019), but with a low atmospheric CO2 concentration of 500 ppm, show no direct conflict with available proxy and paleobotanical data and confirm that under the simulated climatic conditions carbon isotope fractionation represents a robust proxy for atmospheric CO2. To explain the earlier CO2 drop we suggest that early forms of vascular land plants have already strongly influenced weathering. This new perspective on the Devonian questions previous ideas about the climatic conditions and earlier explanations for the Devonian mass extinctions.
The second mass extinction investigated in this thesis is the end-Cretaceous mass extinction (66 million years ago) which differs from the Devonian mass extinctions in terms of the processes involved and the timescale on which the extinctions occurred. In the two studies presented here (Brugger et al., 2017, 2021), we model the climatic effects of the Chicxulub impact, one of the proposed causes of the end-Cretaceous extinction, for the first millennium after the impact. The light-dimming effect of stratospheric sulfate aerosols causes severe cooling, with a decrease of global annual mean surface air temperature of at least 26◦C and a recovery to pre-impact temperatures after more than 30 years. The sudden surface cooling of the ocean induces deep convection which brings nutrients from the deep ocean via upwelling to the surface ocean. Using an ocean biogeochemistry model we explore the combined effect of ocean mixing and iron-rich dust originating from the impactor on the marine biosphere. As soon as light levels have recovered, we find a short, but prominent peak in marine net primary productivity. This newly discovered mechanism could result in toxic effects for marine near-surface ecosystems. Comparison of our model results to proxy data (Vellekoop et al., 2014, 2016, Hull et al., 2020) suggests that carbon release from the terrestrial biosphere is required in addition to the carbon dioxide which can be attributed to the target material. Surface ocean acidification caused by the addition of carbon dioxide and sulfur is only moderate. Taken together, the results indicate a significant contribution of the Chicxulub impact to the end-Cretaceous mass extinction by triggering multiple stressors for the Earth system.
Although the sixth extinction we face today is characterized by human intervention in nature, this thesis shows that we can gain many insights into future extinctions from studying past mass extinctions, such as the importance of the rate of change (Rothman, 2017), the interplay of multiple stressors (Gunderson et al., 2016), and changes in the carbon cycle (Rothman, 2017, Tierney et al., 2020).
The Arctic environments constitute rich and dynamic ecosystems, dominated by microorganisms extremely well adapted to survive and function under severe conditions. A range of physiological adaptations allow the microbiota in these habitats to withstand low temperatures, low water and nutrient availability, high levels of UV radiation, etc. In addition, other adaptations of clear competitive nature are directed at not only surviving but thriving in these environments, by disrupting the metabolism of neighboring cells and affecting intermicrobial communication. Since Arctic microbes are bioindicators which amplify climate alterations in the environment, the Arctic region presents the opportunity to study local microbiota and carry out research about interesting, potentially virulent phenotypes that could be dispersed into other habitats around the globe as a consequence of accelerating climate change. In this context, exploration of Arctic habitats as well as descriptions of the microbes inhabiting them are abundant but microbial competitive strategies commonly associated with virulence and pathogens are rarely reported. In this project, environmental samples from the Arctic region were collected and microorganisms (bacteria and fungi) were isolated. The clinical relevance of these microorganisms was assessed by observing the following virulence markers: ability to grow at a range of temperatures, expression of antimicrobial resistance and production of hemolysins. The aim of this project is to determine the frequency and relevance of these characteristics in an effort to understand microbial adaptations in habitats threatened by climate change. The isolates obtained and described here were able to grow at a range of temperatures, in some cases more than 30 °C higher than their original isolation temperature. A considerable number of them consistently expressed compounds capable of lysing sheep and bovine erythrocytes on blood agar at different incubation temperatures. Ethanolic extracts of these bacteria were able to cause rapid and complete lysis of erythrocyte suspensions and might even be hemolytic when assayed on human blood. In silico analyses showed a variety of resistance elements, some of them novel, against natural and synthetic antimicrobial compounds. In vitro experiments against a number of antimicrobial compounds showed resistance phenotypes belonging to wild-type populations and some non-wild type which clearly denote human influence in the acquisition of antimicrobial resistance. The results of this project demonstrate the presence of virulence-associated factors expressed by microorganisms of natural, non-clinical environments. This study contains some of the first reports, to the best of our knowledge, of hemolytic microbes isolated from the Arctic region. In addition, it provides additional information about the presence and expression of intrinsic and acquired antimicrobial resistance in environmental isolates, contributing to the understanding of the evolution of relevant pathogenic species and opportunistic pathogens. Finally, this study highlights some of the potential risks associated with changes in the polar regions (habitat melting and destruction, ecosystem transition and re-colonization) as important indirect consequences of global warming and altered climatic conditions around the planet.