The return of the mammoth steppe?: Rewilding in northeastern Yakutia and the actual impact of large herbivore grazing on vegetation

Rewilding aims at the restoration of lost ecosystems by re-introducing large herbivores. In northern Siberia, the demise of the mammoth steppe ecosystem at the end of the Pleistocene has been related to the loss of megafauna due to human overhunting. Others argue that climate change at the beginning of the Holocene has triggered the shift from dry, cold steppe vegetation to wet and low productive tundra and taiga vegetation. Despite many different opinions and ongoing discussions on the topic, few case studies are available to test the proposed hypotheses. In this thesis I try to bridge the theoretical backgrounds of palaeoecology and contemporary grazing ecology, and apply these to new data from grazed steppes and surrounding vegetation in Yakutia. This study region is suitable to shed light on the importance of grazers for the (mammoth) steppe vegetation because Yakutia was dominated by mammoth steppe in Pleistocene glacials, and the extrazonal steppes of today are considered potential relics; permafrost deposits in close proximity to these steppes allow palaeobotanical reconstructions of vegetation from the same area; and two game parks, one in Central Yakutia, one in northeastern Yakutia, allow to study grazing impact on contemporary vegetation, specifically steppes. The first part of the thesis focuses on current grassland and steppe vegetation in Russia in general and in Yakutia specifically. Chapter highlights the biological diversity of Russia´s grasslands under diverse climatic and edaphic consditions. It highlights the value of Russian steppes for nature conservation, shows that most grasslands are of agricultural origin and that cessation of land use can pose a threat to both natural and secondary grasslands. Chapter 2 focuses on the phytosociology of extrazonal steppes and other grasslands of Yakutia, and on the harsh climatic and the special edaphic conditions they inhabit. It demonstrates relationships to southern zonal steppes, despite lower species diversity and unique associations with a high contribution of alpine plants. The second part of the thesis addresses the evidence of grazing in the palaorecord as well as effects and importance of grazing for contemporary vegetation. Chapter 3 aims at a comparison of current vegetation with Pleistocene fossil remains in order to find the closest analogues of mammoth steppe vegetation. It demonstrates that meadow steppes formed large parts of the vegetation in both cold and warm stages, only shifting in proportions. Disturbance indicators from grazing animals were more common in the fossil record than today. Chapter 4 discusses the influence of grazing on current vegetation under the given harsh climate; on plant species and trait composition, as well as on vegetation productivity. Climate and soil conditions seem to be the most important determinants of steppe and surrounding vegetation. Large grazers like bison can alter vegetation structure and plant communities on the local scale, but do not alter composition dramatically nor do they increase vegetation productivity in the given settings. In summary, steppes are an important part of biodiversity in Russia and specifically in Yakutia today. The extrazonal steppes of Yakutia are no direct relics of the mammoth steppe, and are not dependent on grazing. However, grazing of large herbivores, such as bison, can locally open up vegetation, and in a continental climate, drive grasslands towards a more steppic character. Further research is needed to investigate the details of these processes and how they could translate to the landscape scale.:Chapter 1: General Introduction 1 1.1. Large mammal extinctions and the onset of the Anthropocene 1 1.2. The 'mammoth steppe' 3 1.2.1. Definitions and concepts 3 1.2.2. Environmental conditions 4 1.2.3. Vegetation of the mammoth steppe 6 1.2.4. Megafauna and suggested ecosystem processes of the mammoth steppe 7 1.2.5. The demise of the mammoth steppe 9 1.3. Grazing ecology 11 1.3.1. Effects of herbivores on vegetation 11 1.3.2. Effects of herbivores on ecosystem processes 14 1.4. Study area 15 1.4.1. Central and northeastern Yakutia 15 1.4.2. Pleistocene Park and the Ust-Buotoma Bisonary 18 1.5. Chapter outline 19 Chapter 2: Land Use of Natural and Secondary Grasslands in Russia 20 2.1. Introduction 21 2.2. Origin and types of grasslands 26 2.2.1. Natural grasslands 26 2.2.1.1. Steppes 26 2.2.1.2. Alpine grasslands 32 2.2.1.3. Azonal grasslands 33 2.2.2. Secondary grasslands 34 2.3. Agronomic Use of grasslands 34 2.3.1. History of land use 34 2.3.2. Current practice of grassland management 38 2.4. Ecological and economic value of grasslands 40 2.5. Threats to grasslands 43 2.6. Conservation of grasslands 48 2.6.1. Legal aspects 48 2.6.2. Prioritization 50 2.6.3. Conservation of grasslands: Protected Areas (PAs) 51 2.7. Conclusions 53 Chapter 3: Extrazonal steppes and other temperate grasslands of northern Siberia - Phytosociological classification and ecological characterization 55 3.1. Introduction 57 3.2. Methods 60 3.3. Results 61 3.3.1. Overview: classification and ordination 61 3.3.2. Description of communities 67 3.3.2.1. Meadows and wet grasslands 67 3.3.2.2. Steppes 68 3.3.3. Phytosociological synopsis 73 3.4. Discussion 75 3.4.1. Meadows 75 3.4.2. Steppes 76 3.4.3. Tundra steppes 80 3.5. Conclusions 81 Cahpter 4: Woodlands and steppes: Pleistocene vegetation in Yakutia´s most continental part recorded in the Batagay permafrost sequence 82 4.1. Introduction 84 4.1.1. Regional setting 86 4.2. Material and methods 87 4.2.1. Material 87 4.2.2. Sampling and preparation 90 4.2.3. Macrofossil preparation and identification 91 4.2.4. Charcoal 91 4.2.5. Palynology preparation and identification 91 4.2.6. Invertebrate sampling and identification 92 4.2.7. Reconstruction of palaeo-vegetation 92 4.3. Results 94 4.3.1. Vegetation of the last cold stage 95 4.3.2. Vegetation of the last interglacial 107 4.3.3. History of local vegetation and environment throughout the sequence 113 4.3.3.1. Unit IV 113 4.3.3.2. Unit III 114 4.3.3.3. Unit II 114 4.4. Discussion 121 4.4.1. Steppes persistent throughout the investigated timespan 121 4.4.2. Steppes in northeast Siberia: Pleistocene survivors or Holocene immigrants? 124 4.4.3. Climatic implications 126 4.5. Conclusions 129 Chapter 5: Grazing at the limit effects of large herbivore grazing on relics of presumed mammoth steppe in NE-Siberia 131 5.1. Introduction 133 5.2. Methods 137 5.2.1. Field sampling and data collection 137 5.2.2. Data analysis 138 5.2.2.1. Species composition 138 5.2.2.2. Plant trait composition 138 5.2.2.3. Taxonomic and functional diversity 139 5.2.2.4. Productivity and chemical composition of vegetation 139 5.3. Results 139 5.3.1. Species composition 140 5.3.2. Trait composition 142 5.3.3. Taxonomic and functional diversity 143 5.3.4. Productivity and chemical composition of vegetation 144 5.4. Discussion 145 5.4.1. Effect of microclimate 145 5.4.2. Grazing effects 146 5.4.3. Implications for rewilding in Siberia 149 Chapter 6: Synthesis - Yakutian steppes and rewilding the mammoth steppe 151 6.1. Contemporary grasslands and herbivore pastures in Yakutia 151 6.2. Contemporary steppes and tundra steppes 152 6.2.1. Relics of the mammoth steppe?! 154 6.3. The role of grazers today - and their proposed role in the mammoth steppe 157 6.4. Outlook on rewilding 166 Chapter 7: Conclusions 174 Chapter 8: Summary 175 Chapter 9: Zusammenfassung 177 II. References 179 III. Appendix 207 IV. List of abbreviations 263 V. Curriculum vitae 270Mit der Auswilderung von Großherbivoren wird oft das Ziel verfolgt, ein lange vergangenes Ökosystem wiederherzustellen. In Nordsibirien ist das die Mammutsteppe, deren Verlust auf die Ausrottung der Großherbivoren-Fauna durch menschliche Überjagung zurückgeführt wurde. Andererseits könnte auch der Klimawandel am Beginn des Holozäns für den Vegetationswandel von trockener, kalter Steppe zu feuchter und wenig produktiver Tundra und Taiga verantwortlich sein. Das Thema wird von verschiedenen Meinungen beherrscht und die Diskussion darüber dauert an; doch konkrete Studien, die diese Hypothesen überprüfen würden, sind selten. In meiner Dissertation versuche ich, eine Brücke zwischen Theorien aus der Paläoökologie und der rezenten Beweidungsökologie zu schlagen und diese auf die beweideten Steppen Yakutiens und deren umgebende Vegetation anzuwenden. Diese Studie liefert Erkenntnisse zur Bedeutung von Weidetieren für die (Mammut-)steppenvegetation, da Yakutien in Pleistozänen Kaltzeiten von Mammutsteppe bedeckt war und die rezenten, extrazonalen Steppen als potenzielle Reliktvegetation angesehen werden; Permafrostaufschlüsse in direkter Umgebung dieser Steppen die paläobotanische Rekonstruktion des selben Gebietes erlauben; und zwei Wildparks, einer in Zentralyakutien, einer im Nordosten Yakutiens, die Erforschung von Beweidungseffekten auf die rezente Vegetation, vor allem Steppenvegetation, ermöglichen. Der erste Teil meiner Dissertation bezieht sich auf die rezente Grasland- und Steppenvegetation Russlands mit Fokus auf Yakutien. Kapitel 1 stellt die biologische Vielfalt der Russischen Grasländer unter verschiedensten klimatischen und edaphischen Bedingungen heraus. Dabei steht die Bedeutung der Russischen Steppen für den Naturschutz im Fokus. Es wird deutlich, dass der Großteil der Grasländer auf landwirtschaftliche Nutzung zurückzuführen ist und dass eine Nutzungsaufgabe sowohl natürliche als auch sekundäre Grasländer bedroht. Kapitel 2 beschäftigt sich mit der Pflanzensozologie der extrazonalen Steppen und anderer Grasländer Yakutiens, sowie mit den speziellen klimatischen und edaphischen Bedingungen, unter denen sie existieren. Ich zeige die verwandtschaftlichen Beziehungen dieser Steppen mit den südlichen, zonalen Steppen, trotz ihrer geringeren Artenvielfalt und ihrer einzigartigen Assoziationen mit hohem Anteil alpiner Arten. Der zweite Teil meiner Dissertation beschäftigt sich mit den Hinweisen auf Beweidung in paläobotanischen Rekonstruktionen der Vegetation, sowie der Bedeutung von Beweidung in der rezenten Vegetation. Kapitel 3 stellt einen Vergleich zwischen der Artenzusammensetzung Pleistozäner Pflanzenfossilien mit denen rezenter Pflanzengesellschaften an, um die Vegetation zu definieren, die der Mammutsteppe am ähnlichsten ist. Dieser Vergleich zeigt, dass Wiesensteppen sowohl in Kalt- als auch in Warmzeiten vorkamen und nur in ihrem Anteil an der Gesamtvegetation schwankten. Störungszeiger für Beweidung waren häufiger in den fossilen Pflanzenresten zu finden als in rezenten Pflanzengesellschaften. Kapitel 4 diskutiert schließlich den Einfluss von Beweidung auf die rezente Vegetation unter den gegebenen extremen Klimaverhältnissen; auf die Zusammensetzung von Arten und Artmerkmalen einer Pflanzengesellschaft, sowie auf deren Produktivität. Die klimatischen und edaphischen Bedingungen scheinen ausschlaggebend für die Steppen- und umgebende Vegetation zu sein. Großherbivoren wie das Bison können allerdings lokal Veränderungen in der Vegetationsstruktur und Pflanzengesellschaften bewirken, wenn auch, unter den gegebenen Umständen, weder Artenzusammensetzung noch Produktivität drastisch verändert wurden. Steppen stellen einen bedeutenden Anteil der Russischen, und besonders Yakutischen, Biodiversität dar. Die extrazonalen Steppen Yakutiens sind keine unmittelbaren Relikte der Mammutsteppe und sind heute nicht auf Beweidung angewiesen. Trotzdem können Großherbivoren wie das Bison lokale Veränderungen bewirken: sie vermögen Baumbestände aufzulichten und, unter kontinentalem Klima, Wiesen hin zu einem mehr steppen-artigen Charakter zu verändern. Weitere Forschung ist nötig, um die Details der beobachteten Prozesse zu erkunden und ihre Übertragbarkeit auf die Landschaftsebene zu überprüfen.:Chapter 1: General Introduction 1 1.1. Large mammal extinctions and the onset of the Anthropocene 1 1.2. The 'mammoth steppe' 3 1.2.1. Definitions and concepts 3 1.2.2. Environmental conditions 4 1.2.3. Vegetation of the mammoth steppe 6 1.2.4. Megafauna and suggested ecosystem processes of the mammoth steppe 7 1.2.5. The demise of the mammoth steppe 9 1.3. Grazing ecology 11 1.3.1. Effects of herbivores on vegetation 11 1.3.2. Effects of herbivores on ecosystem processes 14 1.4. Study area 15 1.4.1. Central and northeastern Yakutia 15 1.4.2. Pleistocene Park and the Ust-Buotoma Bisonary 18 1.5. Chapter outline 19 Chapter 2: Land Use of Natural and Secondary Grasslands in Russia 20 2.1. Introduction 21 2.2. Origin and types of grasslands 26 2.2.1. Natural grasslands 26 2.2.1.1. Steppes 26 2.2.1.2. Alpine grasslands 32 2.2.1.3. Azonal grasslands 33 2.2.2. Secondary grasslands 34 2.3. Agronomic Use of grasslands 34 2.3.1. History of land use 34 2.3.2. Current practice of grassland management 38 2.4. Ecological and economic value of grasslands 40 2.5. Threats to grasslands 43 2.6. Conservation of grasslands 48 2.6.1. Legal aspects 48 2.6.2. Prioritization 50 2.6.3. Conservation of grasslands: Protected Areas (PAs) 51 2.7. Conclusions 53 Chapter 3: Extrazonal steppes and other temperate grasslands of northern Siberia - Phytosociological classification and ecological characterization 55 3.1. Introduction 57 3.2. Methods 60 3.3. Results 61 3.3.1. Overview: classification and ordination 61 3.3.2. Description of communities 67 3.3.2.1. Meadows and wet grasslands 67 3.3.2.2. Steppes 68 3.3.3. Phytosociological synopsis 73 3.4. Discussion 75 3.4.1. Meadows 75 3.4.2. Steppes 76 3.4.3. Tundra steppes 80 3.5. Conclusions 81 Cahpter 4: Woodlands and steppes: Pleistocene vegetation in Yakutia´s most continental part recorded in the Batagay permafrost sequence 82 4.1. Introduction 84 4.1.1. Regional setting 86 4.2. Material and methods 87 4.2.1. Material 87 4.2.2. Sampling and preparation 90 4.2.3. Macrofossil preparation and identification 91 4.2.4. Charcoal 91 4.2.5. Palynology preparation and identification 91 4.2.6. Invertebrate sampling and identification 92 4.2.7. Reconstruction of palaeo-vegetation 92 4.3. Results 94 4.3.1. Vegetation of the last cold stage 95 4.3.2. Vegetation of the last interglacial 107 4.3.3. History of local vegetation and environment throughout the sequence 113 4.3.3.1. Unit IV 113 4.3.3.2. Unit III 114 4.3.3.3. Unit II 114 4.4. Discussion 121 4.4.1. Steppes persistent throughout the investigated timespan 121 4.4.2. Steppes in northeast Siberia: Pleistocene survivors or Holocene immigrants? 124 4.4.3. Climatic implications 126 4.5. Conclusions 129 Chapter 5: Grazing at the limit effects of large herbivore grazing on relics of presumed mammoth steppe in NE-Siberia 131 5.1. Introduction 133 5.2. Methods 137 5.2.1. Field sampling and data collection 137 5.2.2. Data analysis 138 5.2.2.1. Species composition 138 5.2.2.2. Plant trait composition 138 5.2.2.3. Taxonomic and functional diversity 139 5.2.2.4. Productivity and chemical composition of vegetation 139 5.3. Results 139 5.3.1. Species composition 140 5.3.2. Trait composition 142 5.3.3. Taxonomic and functional diversity 143 5.3.4. Productivity and chemical composition of vegetation 144 5.4. Discussion 145 5.4.1. Effect of microclimate 145 5.4.2. Grazing effects 146 5.4.3. Implications for rewilding in Siberia 149 Chapter 6: Synthesis - Yakutian steppes and rewilding the mammoth steppe 151 6.1. Contemporary grasslands and herbivore pastures in Yakutia 151 6.2. Contemporary steppes and tundra steppes 152 6.2.1. Relics of the mammoth steppe?! 154 6.3. The role of grazers today - and their proposed role in the mammoth steppe 157 6.4. Outlook on rewilding 166 Chapter 7: Conclusions 174 Chapter 8: Summary 175 Chapter 9: Zusammenfassung 177 II. References 179 III. Appendix 207 IV. List of abbreviations 263 V. Curriculum vitae 27

Remission and recovery in the Treatment for Adolescents with Depression Study (TADS): acute and long-term outcomes.

OBJECTIVE: We examine remission rate probabilities, recovery rates, and residual symptoms across 36 weeks in the Treatment for Adolescents with Depression Study (TADS). METHOD: The TADS, a multisite clinical trial, randomized 439 adolescents with major depressive disorder to 12 weeks of treatment with fluoxetine, cognitive-behavioral therapy, their combination, or pill placebo. The pill placebo group, treated openly after week 12, was not included in the subsequent analyses. Treatment differences in remission rates and probabilities of remission over time are compared. Recovery rates in remitters at weeks 12 (acute phase remitters) and 18 (continuation phase remitters) are summarized. We also examined whether residual symptoms at the end of 12 weeks of acute treatment predicted later remission. RESULTS: At week 36, the estimated remission rates for intention-to-treat cases were as follows: combination, 60%; fluoxetine, 55%; cognitive-behavioral therapy, 64%; and overall, 60%. Paired comparisons reveal that, at week 24, all active treatments converge on remission outcomes. The recovery rate at week 36 was 65% for acute phase remitters and 71% for continuation phase remitters, with no significant between-treatment differences in recovery rates. Residual symptoms at the end of acute treatment predicted failure to achieve remission at weeks 18 and 36. CONCLUSIONS: Most depressed adolescents in all three treatment modalities achieved remission at the end of 9 months of treatment

Statistics, lessons learned and recommendations from analysis of HIAD 2.0 database

The manuscript firstly describes the data collection and validation process for the European Hydrogen Incidents and Accidents Database (HIAD 2.0), a public repository tool collecting systematic data on hydrogen-related incidents and near-misses. This is followed by an overview of HIAD 2.0, which currently contains 706 events. Subsequently, the approaches and procedures followed by the authors to derive lessons learned and formulate recommendations from the events are described. The lessons learned have been divided into four categories including system design; system manufacturing, installation and modification; human factors and emergency response. An overarching lesson learned is that minor events which occurred simultaneously could still result in serious consequences, echoing James Reason's Swiss Cheese theory. Recommendations were formulated in relation to the established safety principles adapted for hydrogen by the European Hydrogen Safety Panel, considering operational modes, industrial sectors, and human factors. This workprovide an important contribution to the safety of systems involving hydrogen, benefitting technical safety engineers, emergency responders and emergency services. The lesson learned and the discussion derived from the statistics can also be used in training and risk assessment studies, being of equal importance to promote and assist the development of sound safety culture in organisations

Prime Focus Spectrograph (PFS) for the Subaru Telescope: Overview, recent progress, and future perspectives

PFS (Prime Focus Spectrograph), a next generation facility instrument on the 8.2-meter Subaru Telescope, is a very wide-field, massively multiplexed, optical and near-infrared spectrograph. Exploiting the Subaru prime focus, 2394 reconfigurable fibers will be distributed over the 1.3 deg field of view. The spectrograph has been designed with 3 arms of blue, red, and near-infrared cameras to simultaneously observe spectra from 380nm to 1260nm in one exposure at a resolution of ~1.6-2.7A. An international collaboration is developing this instrument under the initiative of Kavli IPMU. The project is now going into the construction phase aiming at undertaking system integration in 2017-2018 and subsequently carrying out engineering operations in 2018-2019. This article gives an overview of the instrument, current project status and future paths forward.Comment: 17 pages, 10 figures. Proceeding of SPIE Astronomical Telescopes and Instrumentation 201

ENIGMA-anxiety working group : Rationale for and organization of large-scale neuroimaging studies of anxiety disorders

Altres ajuts: Anxiety Disorders Research Network European College of Neuropsychopharmacology; Claude Leon Postdoctoral Fellowship; Deutsche Forschungsgemeinschaft (DFG, German Research Foundation, 44541416-TRR58); EU7th Frame Work Marie Curie Actions International Staff Exchange Scheme grant 'European and South African Research Network in Anxiety Disorders' (EUSARNAD); Geestkracht programme of the Netherlands Organization for Health Research and Development (ZonMw, 10-000-1002); Intramural Research Training Award (IRTA) program within the National Institute of Mental Health under the Intramural Research Program (NIMH-IRP, MH002781); National Institute of Mental Health under the Intramural Research Program (NIMH-IRP, ZIA-MH-002782); SA Medical Research Council; U.S. National Institutes of Health grants (P01 AG026572, P01 AG055367, P41 EB015922, R01 AG060610, R56 AG058854, RF1 AG051710, U54 EB020403).Anxiety disorders are highly prevalent and disabling but seem particularly tractable to investigation with translational neuroscience methodologies. Neuroimaging has informed our understanding of the neurobiology of anxiety disorders, but research has been limited by small sample sizes and low statistical power, as well as heterogenous imaging methodology. The ENIGMA-Anxiety Working Group has brought together researchers from around the world, in a harmonized and coordinated effort to address these challenges and generate more robust and reproducible findings. This paper elaborates on the concepts and methods informing the work of the working group to date, and describes the initial approach of the four subgroups studying generalized anxiety disorder, panic disorder, social anxiety disorder, and specific phobia. At present, the ENIGMA-Anxiety database contains information about more than 100 unique samples, from 16 countries and 59 institutes. Future directions include examining additional imaging modalities, integrating imaging and genetic data, and collaborating with other ENIGMA working groups. The ENIGMA consortium creates synergy at the intersection of global mental health and clinical neuroscience, and the ENIGMA-Anxiety Working Group extends the promise of this approach to neuroimaging research on anxiety disorders

GrassPlot - a database of multi-scale plant diversity in Palaearctic grasslands

GrassPlot is a collaborative vegetation-plot database organised by the Eurasian Dry Grassland Group (EDGG) and listed in the Global Index of Vegetation-Plot Databases (GIVD ID EU-00-003). GrassPlot collects plot records (releves) from grasslands and other open habitats of the Palaearctic biogeographic realm. It focuses on precisely delimited plots of eight standard grain sizes (0.0001; 0.001;... 1,000 m(2)) and on nested-plot series with at least four different grain sizes. The usage of GrassPlot is regulated through Bylaws that intend to balance the interests of data contributors and data users. The current version (v. 1.00) contains data for approximately 170,000 plots of different sizes and 2,800 nested-plot series. The key components are richness data and metadata. However, most included datasets also encompass compositional data. About 14,000 plots have near-complete records of terricolous bryophytes and lichens in addition to vascular plants. At present, GrassPlot contains data from 36 countries throughout the Palaearctic, spread across elevational gradients and major grassland types. GrassPlot with its multi-scale and multi-taxon focus complements the larger international vegetationplot databases, such as the European Vegetation Archive (EVA) and the global database " sPlot". Its main aim is to facilitate studies on the scale-and taxon-dependency of biodiversity patterns and drivers along macroecological gradients. GrassPlot is a dynamic database and will expand through new data collection coordinated by the elected Governing Board. We invite researchers with suitable data to join GrassPlot. Researchers with project ideas addressable with GrassPlot data are welcome to submit proposals to the Governing Board

The return of the mammoth steppe?: Rewilding in northeastern Yakutia and the actual impact of large herbivore grazing on vegetation

Rewilding aims at the restoration of lost ecosystems by re-introducing large herbivores. In northern Siberia, the demise of the mammoth steppe ecosystem at the end of the Pleistocene has been related to the loss of megafauna due to human overhunting. Others argue that climate change at the beginning of the Holocene has triggered the shift from dry, cold steppe vegetation to wet and low productive tundra and taiga vegetation. Despite many different opinions and ongoing discussions on the topic, few case studies are available to test the proposed hypotheses. In this thesis I try to bridge the theoretical backgrounds of palaeoecology and contemporary grazing ecology, and apply these to new data from grazed steppes and surrounding vegetation in Yakutia. This study region is suitable to shed light on the importance of grazers for the (mammoth) steppe vegetation because Yakutia was dominated by mammoth steppe in Pleistocene glacials, and the extrazonal steppes of today are considered potential relics; permafrost deposits in close proximity to these steppes allow palaeobotanical reconstructions of vegetation from the same area; and two game parks, one in Central Yakutia, one in northeastern Yakutia, allow to study grazing impact on contemporary vegetation, specifically steppes. The first part of the thesis focuses on current grassland and steppe vegetation in Russia in general and in Yakutia specifically. Chapter highlights the biological diversity of Russia´s grasslands under diverse climatic and edaphic consditions. It highlights the value of Russian steppes for nature conservation, shows that most grasslands are of agricultural origin and that cessation of land use can pose a threat to both natural and secondary grasslands. Chapter 2 focuses on the phytosociology of extrazonal steppes and other grasslands of Yakutia, and on the harsh climatic and the special edaphic conditions they inhabit. It demonstrates relationships to southern zonal steppes, despite lower species diversity and unique associations with a high contribution of alpine plants. The second part of the thesis addresses the evidence of grazing in the palaorecord as well as effects and importance of grazing for contemporary vegetation. Chapter 3 aims at a comparison of current vegetation with Pleistocene fossil remains in order to find the closest analogues of mammoth steppe vegetation. It demonstrates that meadow steppes formed large parts of the vegetation in both cold and warm stages, only shifting in proportions. Disturbance indicators from grazing animals were more common in the fossil record than today. Chapter 4 discusses the influence of grazing on current vegetation under the given harsh climate; on plant species and trait composition, as well as on vegetation productivity. Climate and soil conditions seem to be the most important determinants of steppe and surrounding vegetation. Large grazers like bison can alter vegetation structure and plant communities on the local scale, but do not alter composition dramatically nor do they increase vegetation productivity in the given settings. In summary, steppes are an important part of biodiversity in Russia and specifically in Yakutia today. The extrazonal steppes of Yakutia are no direct relics of the mammoth steppe, and are not dependent on grazing. However, grazing of large herbivores, such as bison, can locally open up vegetation, and in a continental climate, drive grasslands towards a more steppic character. Further research is needed to investigate the details of these processes and how they could translate to the landscape scale.:Chapter 1: General Introduction 1 1.1. Large mammal extinctions and the onset of the Anthropocene 1 1.2. The 'mammoth steppe' 3 1.2.1. Definitions and concepts 3 1.2.2. Environmental conditions 4 1.2.3. Vegetation of the mammoth steppe 6 1.2.4. Megafauna and suggested ecosystem processes of the mammoth steppe 7 1.2.5. The demise of the mammoth steppe 9 1.3. Grazing ecology 11 1.3.1. Effects of herbivores on vegetation 11 1.3.2. Effects of herbivores on ecosystem processes 14 1.4. Study area 15 1.4.1. Central and northeastern Yakutia 15 1.4.2. Pleistocene Park and the Ust-Buotoma Bisonary 18 1.5. Chapter outline 19 Chapter 2: Land Use of Natural and Secondary Grasslands in Russia 20 2.1. Introduction 21 2.2. Origin and types of grasslands 26 2.2.1. Natural grasslands 26 2.2.1.1. Steppes 26 2.2.1.2. Alpine grasslands 32 2.2.1.3. Azonal grasslands 33 2.2.2. Secondary grasslands 34 2.3. Agronomic Use of grasslands 34 2.3.1. History of land use 34 2.3.2. Current practice of grassland management 38 2.4. Ecological and economic value of grasslands 40 2.5. Threats to grasslands 43 2.6. Conservation of grasslands 48 2.6.1. Legal aspects 48 2.6.2. Prioritization 50 2.6.3. Conservation of grasslands: Protected Areas (PAs) 51 2.7. Conclusions 53 Chapter 3: Extrazonal steppes and other temperate grasslands of northern Siberia - Phytosociological classification and ecological characterization 55 3.1. Introduction 57 3.2. Methods 60 3.3. Results 61 3.3.1. Overview: classification and ordination 61 3.3.2. Description of communities 67 3.3.2.1. Meadows and wet grasslands 67 3.3.2.2. Steppes 68 3.3.3. Phytosociological synopsis 73 3.4. Discussion 75 3.4.1. Meadows 75 3.4.2. Steppes 76 3.4.3. Tundra steppes 80 3.5. Conclusions 81 Cahpter 4: Woodlands and steppes: Pleistocene vegetation in Yakutia´s most continental part recorded in the Batagay permafrost sequence 82 4.1. Introduction 84 4.1.1. Regional setting 86 4.2. Material and methods 87 4.2.1. Material 87 4.2.2. Sampling and preparation 90 4.2.3. Macrofossil preparation and identification 91 4.2.4. Charcoal 91 4.2.5. Palynology preparation and identification 91 4.2.6. Invertebrate sampling and identification 92 4.2.7. Reconstruction of palaeo-vegetation 92 4.3. Results 94 4.3.1. Vegetation of the last cold stage 95 4.3.2. Vegetation of the last interglacial 107 4.3.3. History of local vegetation and environment throughout the sequence 113 4.3.3.1. Unit IV 113 4.3.3.2. Unit III 114 4.3.3.3. Unit II 114 4.4. Discussion 121 4.4.1. Steppes persistent throughout the investigated timespan 121 4.4.2. Steppes in northeast Siberia: Pleistocene survivors or Holocene immigrants? 124 4.4.3. Climatic implications 126 4.5. Conclusions 129 Chapter 5: Grazing at the limit effects of large herbivore grazing on relics of presumed mammoth steppe in NE-Siberia 131 5.1. Introduction 133 5.2. Methods 137 5.2.1. Field sampling and data collection 137 5.2.2. Data analysis 138 5.2.2.1. Species composition 138 5.2.2.2. Plant trait composition 138 5.2.2.3. Taxonomic and functional diversity 139 5.2.2.4. Productivity and chemical composition of vegetation 139 5.3. Results 139 5.3.1. Species composition 140 5.3.2. Trait composition 142 5.3.3. Taxonomic and functional diversity 143 5.3.4. Productivity and chemical composition of vegetation 144 5.4. Discussion 145 5.4.1. Effect of microclimate 145 5.4.2. Grazing effects 146 5.4.3. Implications for rewilding in Siberia 149 Chapter 6: Synthesis - Yakutian steppes and rewilding the mammoth steppe 151 6.1. Contemporary grasslands and herbivore pastures in Yakutia 151 6.2. Contemporary steppes and tundra steppes 152 6.2.1. Relics of the mammoth steppe?! 154 6.3. The role of grazers today - and their proposed role in the mammoth steppe 157 6.4. Outlook on rewilding 166 Chapter 7: Conclusions 174 Chapter 8: Summary 175 Chapter 9: Zusammenfassung 177 II. References 179 III. Appendix 207 IV. List of abbreviations 263 V. Curriculum vitae 270Mit der Auswilderung von Großherbivoren wird oft das Ziel verfolgt, ein lange vergangenes Ökosystem wiederherzustellen. In Nordsibirien ist das die Mammutsteppe, deren Verlust auf die Ausrottung der Großherbivoren-Fauna durch menschliche Überjagung zurückgeführt wurde. Andererseits könnte auch der Klimawandel am Beginn des Holozäns für den Vegetationswandel von trockener, kalter Steppe zu feuchter und wenig produktiver Tundra und Taiga verantwortlich sein. Das Thema wird von verschiedenen Meinungen beherrscht und die Diskussion darüber dauert an; doch konkrete Studien, die diese Hypothesen überprüfen würden, sind selten. In meiner Dissertation versuche ich, eine Brücke zwischen Theorien aus der Paläoökologie und der rezenten Beweidungsökologie zu schlagen und diese auf die beweideten Steppen Yakutiens und deren umgebende Vegetation anzuwenden. Diese Studie liefert Erkenntnisse zur Bedeutung von Weidetieren für die (Mammut-)steppenvegetation, da Yakutien in Pleistozänen Kaltzeiten von Mammutsteppe bedeckt war und die rezenten, extrazonalen Steppen als potenzielle Reliktvegetation angesehen werden; Permafrostaufschlüsse in direkter Umgebung dieser Steppen die paläobotanische Rekonstruktion des selben Gebietes erlauben; und zwei Wildparks, einer in Zentralyakutien, einer im Nordosten Yakutiens, die Erforschung von Beweidungseffekten auf die rezente Vegetation, vor allem Steppenvegetation, ermöglichen. Der erste Teil meiner Dissertation bezieht sich auf die rezente Grasland- und Steppenvegetation Russlands mit Fokus auf Yakutien. Kapitel 1 stellt die biologische Vielfalt der Russischen Grasländer unter verschiedensten klimatischen und edaphischen Bedingungen heraus. Dabei steht die Bedeutung der Russischen Steppen für den Naturschutz im Fokus. Es wird deutlich, dass der Großteil der Grasländer auf landwirtschaftliche Nutzung zurückzuführen ist und dass eine Nutzungsaufgabe sowohl natürliche als auch sekundäre Grasländer bedroht. Kapitel 2 beschäftigt sich mit der Pflanzensozologie der extrazonalen Steppen und anderer Grasländer Yakutiens, sowie mit den speziellen klimatischen und edaphischen Bedingungen, unter denen sie existieren. Ich zeige die verwandtschaftlichen Beziehungen dieser Steppen mit den südlichen, zonalen Steppen, trotz ihrer geringeren Artenvielfalt und ihrer einzigartigen Assoziationen mit hohem Anteil alpiner Arten. Der zweite Teil meiner Dissertation beschäftigt sich mit den Hinweisen auf Beweidung in paläobotanischen Rekonstruktionen der Vegetation, sowie der Bedeutung von Beweidung in der rezenten Vegetation. Kapitel 3 stellt einen Vergleich zwischen der Artenzusammensetzung Pleistozäner Pflanzenfossilien mit denen rezenter Pflanzengesellschaften an, um die Vegetation zu definieren, die der Mammutsteppe am ähnlichsten ist. Dieser Vergleich zeigt, dass Wiesensteppen sowohl in Kalt- als auch in Warmzeiten vorkamen und nur in ihrem Anteil an der Gesamtvegetation schwankten. Störungszeiger für Beweidung waren häufiger in den fossilen Pflanzenresten zu finden als in rezenten Pflanzengesellschaften. Kapitel 4 diskutiert schließlich den Einfluss von Beweidung auf die rezente Vegetation unter den gegebenen extremen Klimaverhältnissen; auf die Zusammensetzung von Arten und Artmerkmalen einer Pflanzengesellschaft, sowie auf deren Produktivität. Die klimatischen und edaphischen Bedingungen scheinen ausschlaggebend für die Steppen- und umgebende Vegetation zu sein. Großherbivoren wie das Bison können allerdings lokal Veränderungen in der Vegetationsstruktur und Pflanzengesellschaften bewirken, wenn auch, unter den gegebenen Umständen, weder Artenzusammensetzung noch Produktivität drastisch verändert wurden. Steppen stellen einen bedeutenden Anteil der Russischen, und besonders Yakutischen, Biodiversität dar. Die extrazonalen Steppen Yakutiens sind keine unmittelbaren Relikte der Mammutsteppe und sind heute nicht auf Beweidung angewiesen. Trotzdem können Großherbivoren wie das Bison lokale Veränderungen bewirken: sie vermögen Baumbestände aufzulichten und, unter kontinentalem Klima, Wiesen hin zu einem mehr steppen-artigen Charakter zu verändern. Weitere Forschung ist nötig, um die Details der beobachteten Prozesse zu erkunden und ihre Übertragbarkeit auf die Landschaftsebene zu überprüfen.:Chapter 1: General Introduction 1 1.1. Large mammal extinctions and the onset of the Anthropocene 1 1.2. The 'mammoth steppe' 3 1.2.1. Definitions and concepts 3 1.2.2. Environmental conditions 4 1.2.3. Vegetation of the mammoth steppe 6 1.2.4. Megafauna and suggested ecosystem processes of the mammoth steppe 7 1.2.5. The demise of the mammoth steppe 9 1.3. Grazing ecology 11 1.3.1. Effects of herbivores on vegetation 11 1.3.2. Effects of herbivores on ecosystem processes 14 1.4. Study area 15 1.4.1. Central and northeastern Yakutia 15 1.4.2. Pleistocene Park and the Ust-Buotoma Bisonary 18 1.5. Chapter outline 19 Chapter 2: Land Use of Natural and Secondary Grasslands in Russia 20 2.1. Introduction 21 2.2. Origin and types of grasslands 26 2.2.1. Natural grasslands 26 2.2.1.1. Steppes 26 2.2.1.2. Alpine grasslands 32 2.2.1.3. Azonal grasslands 33 2.2.2. Secondary grasslands 34 2.3. Agronomic Use of grasslands 34 2.3.1. History of land use 34 2.3.2. Current practice of grassland management 38 2.4. Ecological and economic value of grasslands 40 2.5. Threats to grasslands 43 2.6. Conservation of grasslands 48 2.6.1. Legal aspects 48 2.6.2. Prioritization 50 2.6.3. Conservation of grasslands: Protected Areas (PAs) 51 2.7. Conclusions 53 Chapter 3: Extrazonal steppes and other temperate grasslands of northern Siberia - Phytosociological classification and ecological characterization 55 3.1. Introduction 57 3.2. Methods 60 3.3. Results 61 3.3.1. Overview: classification and ordination 61 3.3.2. Description of communities 67 3.3.2.1. Meadows and wet grasslands 67 3.3.2.2. Steppes 68 3.3.3. Phytosociological synopsis 73 3.4. Discussion 75 3.4.1. Meadows 75 3.4.2. Steppes 76 3.4.3. Tundra steppes 80 3.5. Conclusions 81 Cahpter 4: Woodlands and steppes: Pleistocene vegetation in Yakutia´s most continental part recorded in the Batagay permafrost sequence 82 4.1. Introduction 84 4.1.1. Regional setting 86 4.2. Material and methods 87 4.2.1. Material 87 4.2.2. Sampling and preparation 90 4.2.3. Macrofossil preparation and identification 91 4.2.4. Charcoal 91 4.2.5. Palynology preparation and identification 91 4.2.6. Invertebrate sampling and identification 92 4.2.7. Reconstruction of palaeo-vegetation 92 4.3. Results 94 4.3.1. Vegetation of the last cold stage 95 4.3.2. Vegetation of the last interglacial 107 4.3.3. History of local vegetation and environment throughout the sequence 113 4.3.3.1. Unit IV 113 4.3.3.2. Unit III 114 4.3.3.3. Unit II 114 4.4. Discussion 121 4.4.1. Steppes persistent throughout the investigated timespan 121 4.4.2. Steppes in northeast Siberia: Pleistocene survivors or Holocene immigrants? 124 4.4.3. Climatic implications 126 4.5. Conclusions 129 Chapter 5: Grazing at the limit effects of large herbivore grazing on relics of presumed mammoth steppe in NE-Siberia 131 5.1. Introduction 133 5.2. Methods 137 5.2.1. Field sampling and data collection 137 5.2.2. Data analysis 138 5.2.2.1. Species composition 138 5.2.2.2. Plant trait composition 138 5.2.2.3. Taxonomic and functional diversity 139 5.2.2.4. Productivity and chemical composition of vegetation 139 5.3. Results 139 5.3.1. Species composition 140 5.3.2. Trait composition 142 5.3.3. Taxonomic and functional diversity 143 5.3.4. Productivity and chemical composition of vegetation 144 5.4. Discussion 145 5.4.1. Effect of microclimate 145 5.4.2. Grazing effects 146 5.4.3. Implications for rewilding in Siberia 149 Chapter 6: Synthesis - Yakutian steppes and rewilding the mammoth steppe 151 6.1. Contemporary grasslands and herbivore pastures in Yakutia 151 6.2. Contemporary steppes and tundra steppes 152 6.2.1. Relics of the mammoth steppe?! 154 6.3. The role of grazers today - and their proposed role in the mammoth steppe 157 6.4. Outlook on rewilding 166 Chapter 7: Conclusions 174 Chapter 8: Summary 175 Chapter 9: Zusammenfassung 177 II. References 179 III. Appendix 207 IV. List of abbreviations 263 V. Curriculum vitae 27

The return of the mammoth steppe?: Rewilding in northeastern Yakutia and the actual impact of large herbivore grazing on vegetation

Rewilding aims at the restoration of lost ecosystems by re-introducing large herbivores. In northern Siberia, the demise of the mammoth steppe ecosystem at the end of the Pleistocene has been related to the loss of megafauna due to human overhunting. Others argue that climate change at the beginning of the Holocene has triggered the shift from dry, cold steppe vegetation to wet and low productive tundra and taiga vegetation. Despite many different opinions and ongoing discussions on the topic, few case studies are available to test the proposed hypotheses. In this thesis I try to bridge the theoretical backgrounds of palaeoecology and contemporary grazing ecology, and apply these to new data from grazed steppes and surrounding vegetation in Yakutia. This study region is suitable to shed light on the importance of grazers for the (mammoth) steppe vegetation because Yakutia was dominated by mammoth steppe in Pleistocene glacials, and the extrazonal steppes of today are considered potential relics; permafrost deposits in close proximity to these steppes allow palaeobotanical reconstructions of vegetation from the same area; and two game parks, one in Central Yakutia, one in northeastern Yakutia, allow to study grazing impact on contemporary vegetation, specifically steppes. The first part of the thesis focuses on current grassland and steppe vegetation in Russia in general and in Yakutia specifically. Chapter highlights the biological diversity of Russia´s grasslands under diverse climatic and edaphic consditions. It highlights the value of Russian steppes for nature conservation, shows that most grasslands are of agricultural origin and that cessation of land use can pose a threat to both natural and secondary grasslands. Chapter 2 focuses on the phytosociology of extrazonal steppes and other grasslands of Yakutia, and on the harsh climatic and the special edaphic conditions they inhabit. It demonstrates relationships to southern zonal steppes, despite lower species diversity and unique associations with a high contribution of alpine plants. The second part of the thesis addresses the evidence of grazing in the palaorecord as well as effects and importance of grazing for contemporary vegetation. Chapter 3 aims at a comparison of current vegetation with Pleistocene fossil remains in order to find the closest analogues of mammoth steppe vegetation. It demonstrates that meadow steppes formed large parts of the vegetation in both cold and warm stages, only shifting in proportions. Disturbance indicators from grazing animals were more common in the fossil record than today. Chapter 4 discusses the influence of grazing on current vegetation under the given harsh climate; on plant species and trait composition, as well as on vegetation productivity. Climate and soil conditions seem to be the most important determinants of steppe and surrounding vegetation. Large grazers like bison can alter vegetation structure and plant communities on the local scale, but do not alter composition dramatically nor do they increase vegetation productivity in the given settings. In summary, steppes are an important part of biodiversity in Russia and specifically in Yakutia today. The extrazonal steppes of Yakutia are no direct relics of the mammoth steppe, and are not dependent on grazing. However, grazing of large herbivores, such as bison, can locally open up vegetation, and in a continental climate, drive grasslands towards a more steppic character. Further research is needed to investigate the details of these processes and how they could translate to the landscape scale.:Chapter 1: General Introduction 1 1.1. Large mammal extinctions and the onset of the Anthropocene 1 1.2. The 'mammoth steppe' 3 1.2.1. Definitions and concepts 3 1.2.2. Environmental conditions 4 1.2.3. Vegetation of the mammoth steppe 6 1.2.4. Megafauna and suggested ecosystem processes of the mammoth steppe 7 1.2.5. The demise of the mammoth steppe 9 1.3. Grazing ecology 11 1.3.1. Effects of herbivores on vegetation 11 1.3.2. Effects of herbivores on ecosystem processes 14 1.4. Study area 15 1.4.1. Central and northeastern Yakutia 15 1.4.2. Pleistocene Park and the Ust-Buotoma Bisonary 18 1.5. Chapter outline 19 Chapter 2: Land Use of Natural and Secondary Grasslands in Russia 20 2.1. Introduction 21 2.2. Origin and types of grasslands 26 2.2.1. Natural grasslands 26 2.2.1.1. Steppes 26 2.2.1.2. Alpine grasslands 32 2.2.1.3. Azonal grasslands 33 2.2.2. Secondary grasslands 34 2.3. Agronomic Use of grasslands 34 2.3.1. History of land use 34 2.3.2. Current practice of grassland management 38 2.4. Ecological and economic value of grasslands 40 2.5. Threats to grasslands 43 2.6. Conservation of grasslands 48 2.6.1. Legal aspects 48 2.6.2. Prioritization 50 2.6.3. Conservation of grasslands: Protected Areas (PAs) 51 2.7. Conclusions 53 Chapter 3: Extrazonal steppes and other temperate grasslands of northern Siberia - Phytosociological classification and ecological characterization 55 3.1. Introduction 57 3.2. Methods 60 3.3. Results 61 3.3.1. Overview: classification and ordination 61 3.3.2. Description of communities 67 3.3.2.1. Meadows and wet grasslands 67 3.3.2.2. Steppes 68 3.3.3. Phytosociological synopsis 73 3.4. Discussion 75 3.4.1. Meadows 75 3.4.2. Steppes 76 3.4.3. Tundra steppes 80 3.5. Conclusions 81 Cahpter 4: Woodlands and steppes: Pleistocene vegetation in Yakutia´s most continental part recorded in the Batagay permafrost sequence 82 4.1. Introduction 84 4.1.1. Regional setting 86 4.2. Material and methods 87 4.2.1. Material 87 4.2.2. Sampling and preparation 90 4.2.3. Macrofossil preparation and identification 91 4.2.4. Charcoal 91 4.2.5. Palynology preparation and identification 91 4.2.6. Invertebrate sampling and identification 92 4.2.7. Reconstruction of palaeo-vegetation 92 4.3. Results 94 4.3.1. Vegetation of the last cold stage 95 4.3.2. Vegetation of the last interglacial 107 4.3.3. History of local vegetation and environment throughout the sequence 113 4.3.3.1. Unit IV 113 4.3.3.2. Unit III 114 4.3.3.3. Unit II 114 4.4. Discussion 121 4.4.1. Steppes persistent throughout the investigated timespan 121 4.4.2. Steppes in northeast Siberia: Pleistocene survivors or Holocene immigrants? 124 4.4.3. Climatic implications 126 4.5. Conclusions 129 Chapter 5: Grazing at the limit effects of large herbivore grazing on relics of presumed mammoth steppe in NE-Siberia 131 5.1. Introduction 133 5.2. Methods 137 5.2.1. Field sampling and data collection 137 5.2.2. Data analysis 138 5.2.2.1. Species composition 138 5.2.2.2. Plant trait composition 138 5.2.2.3. Taxonomic and functional diversity 139 5.2.2.4. Productivity and chemical composition of vegetation 139 5.3. Results 139 5.3.1. Species composition 140 5.3.2. Trait composition 142 5.3.3. Taxonomic and functional diversity 143 5.3.4. Productivity and chemical composition of vegetation 144 5.4. Discussion 145 5.4.1. Effect of microclimate 145 5.4.2. Grazing effects 146 5.4.3. Implications for rewilding in Siberia 149 Chapter 6: Synthesis - Yakutian steppes and rewilding the mammoth steppe 151 6.1. Contemporary grasslands and herbivore pastures in Yakutia 151 6.2. Contemporary steppes and tundra steppes 152 6.2.1. Relics of the mammoth steppe?! 154 6.3. The role of grazers today - and their proposed role in the mammoth steppe 157 6.4. Outlook on rewilding 166 Chapter 7: Conclusions 174 Chapter 8: Summary 175 Chapter 9: Zusammenfassung 177 II. References 179 III. Appendix 207 IV. List of abbreviations 263 V. Curriculum vitae 270Mit der Auswilderung von Großherbivoren wird oft das Ziel verfolgt, ein lange vergangenes Ökosystem wiederherzustellen. In Nordsibirien ist das die Mammutsteppe, deren Verlust auf die Ausrottung der Großherbivoren-Fauna durch menschliche Überjagung zurückgeführt wurde. Andererseits könnte auch der Klimawandel am Beginn des Holozäns für den Vegetationswandel von trockener, kalter Steppe zu feuchter und wenig produktiver Tundra und Taiga verantwortlich sein. Das Thema wird von verschiedenen Meinungen beherrscht und die Diskussion darüber dauert an; doch konkrete Studien, die diese Hypothesen überprüfen würden, sind selten. In meiner Dissertation versuche ich, eine Brücke zwischen Theorien aus der Paläoökologie und der rezenten Beweidungsökologie zu schlagen und diese auf die beweideten Steppen Yakutiens und deren umgebende Vegetation anzuwenden. Diese Studie liefert Erkenntnisse zur Bedeutung von Weidetieren für die (Mammut-)steppenvegetation, da Yakutien in Pleistozänen Kaltzeiten von Mammutsteppe bedeckt war und die rezenten, extrazonalen Steppen als potenzielle Reliktvegetation angesehen werden; Permafrostaufschlüsse in direkter Umgebung dieser Steppen die paläobotanische Rekonstruktion des selben Gebietes erlauben; und zwei Wildparks, einer in Zentralyakutien, einer im Nordosten Yakutiens, die Erforschung von Beweidungseffekten auf die rezente Vegetation, vor allem Steppenvegetation, ermöglichen. Der erste Teil meiner Dissertation bezieht sich auf die rezente Grasland- und Steppenvegetation Russlands mit Fokus auf Yakutien. Kapitel 1 stellt die biologische Vielfalt der Russischen Grasländer unter verschiedensten klimatischen und edaphischen Bedingungen heraus. Dabei steht die Bedeutung der Russischen Steppen für den Naturschutz im Fokus. Es wird deutlich, dass der Großteil der Grasländer auf landwirtschaftliche Nutzung zurückzuführen ist und dass eine Nutzungsaufgabe sowohl natürliche als auch sekundäre Grasländer bedroht. Kapitel 2 beschäftigt sich mit der Pflanzensozologie der extrazonalen Steppen und anderer Grasländer Yakutiens, sowie mit den speziellen klimatischen und edaphischen Bedingungen, unter denen sie existieren. Ich zeige die verwandtschaftlichen Beziehungen dieser Steppen mit den südlichen, zonalen Steppen, trotz ihrer geringeren Artenvielfalt und ihrer einzigartigen Assoziationen mit hohem Anteil alpiner Arten. Der zweite Teil meiner Dissertation beschäftigt sich mit den Hinweisen auf Beweidung in paläobotanischen Rekonstruktionen der Vegetation, sowie der Bedeutung von Beweidung in der rezenten Vegetation. Kapitel 3 stellt einen Vergleich zwischen der Artenzusammensetzung Pleistozäner Pflanzenfossilien mit denen rezenter Pflanzengesellschaften an, um die Vegetation zu definieren, die der Mammutsteppe am ähnlichsten ist. Dieser Vergleich zeigt, dass Wiesensteppen sowohl in Kalt- als auch in Warmzeiten vorkamen und nur in ihrem Anteil an der Gesamtvegetation schwankten. Störungszeiger für Beweidung waren häufiger in den fossilen Pflanzenresten zu finden als in rezenten Pflanzengesellschaften. Kapitel 4 diskutiert schließlich den Einfluss von Beweidung auf die rezente Vegetation unter den gegebenen extremen Klimaverhältnissen; auf die Zusammensetzung von Arten und Artmerkmalen einer Pflanzengesellschaft, sowie auf deren Produktivität. Die klimatischen und edaphischen Bedingungen scheinen ausschlaggebend für die Steppen- und umgebende Vegetation zu sein. Großherbivoren wie das Bison können allerdings lokal Veränderungen in der Vegetationsstruktur und Pflanzengesellschaften bewirken, wenn auch, unter den gegebenen Umständen, weder Artenzusammensetzung noch Produktivität drastisch verändert wurden. Steppen stellen einen bedeutenden Anteil der Russischen, und besonders Yakutischen, Biodiversität dar. Die extrazonalen Steppen Yakutiens sind keine unmittelbaren Relikte der Mammutsteppe und sind heute nicht auf Beweidung angewiesen. Trotzdem können Großherbivoren wie das Bison lokale Veränderungen bewirken: sie vermögen Baumbestände aufzulichten und, unter kontinentalem Klima, Wiesen hin zu einem mehr steppen-artigen Charakter zu verändern. Weitere Forschung ist nötig, um die Details der beobachteten Prozesse zu erkunden und ihre Übertragbarkeit auf die Landschaftsebene zu überprüfen.:Chapter 1: General Introduction 1 1.1. Large mammal extinctions and the onset of the Anthropocene 1 1.2. The 'mammoth steppe' 3 1.2.1. Definitions and concepts 3 1.2.2. Environmental conditions 4 1.2.3. Vegetation of the mammoth steppe 6 1.2.4. Megafauna and suggested ecosystem processes of the mammoth steppe 7 1.2.5. The demise of the mammoth steppe 9 1.3. Grazing ecology 11 1.3.1. Effects of herbivores on vegetation 11 1.3.2. Effects of herbivores on ecosystem processes 14 1.4. Study area 15 1.4.1. Central and northeastern Yakutia 15 1.4.2. Pleistocene Park and the Ust-Buotoma Bisonary 18 1.5. Chapter outline 19 Chapter 2: Land Use of Natural and Secondary Grasslands in Russia 20 2.1. Introduction 21 2.2. Origin and types of grasslands 26 2.2.1. Natural grasslands 26 2.2.1.1. Steppes 26 2.2.1.2. Alpine grasslands 32 2.2.1.3. Azonal grasslands 33 2.2.2. Secondary grasslands 34 2.3. Agronomic Use of grasslands 34 2.3.1. History of land use 34 2.3.2. Current practice of grassland management 38 2.4. Ecological and economic value of grasslands 40 2.5. Threats to grasslands 43 2.6. Conservation of grasslands 48 2.6.1. Legal aspects 48 2.6.2. Prioritization 50 2.6.3. Conservation of grasslands: Protected Areas (PAs) 51 2.7. Conclusions 53 Chapter 3: Extrazonal steppes and other temperate grasslands of northern Siberia - Phytosociological classification and ecological characterization 55 3.1. Introduction 57 3.2. Methods 60 3.3. Results 61 3.3.1. Overview: classification and ordination 61 3.3.2. Description of communities 67 3.3.2.1. Meadows and wet grasslands 67 3.3.2.2. Steppes 68 3.3.3. Phytosociological synopsis 73 3.4. Discussion 75 3.4.1. Meadows 75 3.4.2. Steppes 76 3.4.3. Tundra steppes 80 3.5. Conclusions 81 Cahpter 4: Woodlands and steppes: Pleistocene vegetation in Yakutia´s most continental part recorded in the Batagay permafrost sequence 82 4.1. Introduction 84 4.1.1. Regional setting 86 4.2. Material and methods 87 4.2.1. Material 87 4.2.2. Sampling and preparation 90 4.2.3. Macrofossil preparation and identification 91 4.2.4. Charcoal 91 4.2.5. Palynology preparation and identification 91 4.2.6. Invertebrate sampling and identification 92 4.2.7. Reconstruction of palaeo-vegetation 92 4.3. Results 94 4.3.1. Vegetation of the last cold stage 95 4.3.2. Vegetation of the last interglacial 107 4.3.3. History of local vegetation and environment throughout the sequence 113 4.3.3.1. Unit IV 113 4.3.3.2. Unit III 114 4.3.3.3. Unit II 114 4.4. Discussion 121 4.4.1. Steppes persistent throughout the investigated timespan 121 4.4.2. Steppes in northeast Siberia: Pleistocene survivors or Holocene immigrants? 124 4.4.3. Climatic implications 126 4.5. Conclusions 129 Chapter 5: Grazing at the limit effects of large herbivore grazing on relics of presumed mammoth steppe in NE-Siberia 131 5.1. Introduction 133 5.2. Methods 137 5.2.1. Field sampling and data collection 137 5.2.2. Data analysis 138 5.2.2.1. Species composition 138 5.2.2.2. Plant trait composition 138 5.2.2.3. Taxonomic and functional diversity 139 5.2.2.4. Productivity and chemical composition of vegetation 139 5.3. Results 139 5.3.1. Species composition 140 5.3.2. Trait composition 142 5.3.3. Taxonomic and functional diversity 143 5.3.4. Productivity and chemical composition of vegetation 144 5.4. Discussion 145 5.4.1. Effect of microclimate 145 5.4.2. Grazing effects 146 5.4.3. Implications for rewilding in Siberia 149 Chapter 6: Synthesis - Yakutian steppes and rewilding the mammoth steppe 151 6.1. Contemporary grasslands and herbivore pastures in Yakutia 151 6.2. Contemporary steppes and tundra steppes 152 6.2.1. Relics of the mammoth steppe?! 154 6.3. The role of grazers today - and their proposed role in the mammoth steppe 157 6.4. Outlook on rewilding 166 Chapter 7: Conclusions 174 Chapter 8: Summary 175 Chapter 9: Zusammenfassung 177 II. References 179 III. Appendix 207 IV. List of abbreviations 263 V. Curriculum vitae 27

Ferritin Overexpression for Noninvasive Magnetic Resonance Imaging–Based Tracking of Stem Cells Transplanted into the Heart

An unmet need in cardiac cell therapy is a noninvasive imaging technique capable of tracking changes in graft size over time and monitoring cell dynamics such as replication and death, factors to which commonly used superparamagnetic nanoparticles are insensitive. Our goal was to explore if overexpression of ferritin, a nontoxic iron-binding protein, can be used for noninvasive magnetic resonance imaging (MRI) of cells transplanted into the infarcted heart. Mouse skeletal myoblasts (C2C12 cells) were engineered to overexpress ferritin. Ferritin overexpression did not interfere with cell viability, proliferation, or differentiation into multinucleated myotubes. Ferritin overexpression caused a 25% decrease in T 2 relaxation time in vitro compared to wild-type cells. Transgenic grafts were detected in vivo 3 weeks after transplantation into infarcted hearts of syngeneic mice as areas of hypointensity caused by iron accumulation in overexpressed ferritin complexes. Graft size evaluation by MRI correlated tighly with histologic measurements ( R 2 = .8). Our studies demonstrated the feasibility of ferritin overexpression in mouse skeletal myoblasts and the successful detection of transgenic cells by MRI in vitro and in vivo after transplantation into the infarcted mouse heart. These experiments lay the groundwork for using the MRI gene reporter ferritin to track stem cells transplanted to the heart