Thesis (Ph. D.)--University of Wyoming, 2007. / Title from PDF title page (viewed on July 15, 2008). Includes bibliographical references.

Thesis (M.S.)--Oregon State University, 2009. / Printout. Includes bibliographical references (leaves 59-67). Also available on the World Wide Web.

La survie des, populations humaines est directement reliée à la préservation de leur environnement, principalement par le maintien des services écosystémiques. Cependant, depuis les débuts de l'industrialisation, plusieurs actions anthropiques ont fortement modifié les écosystèmes et les services écosystémiques en découlant, sur une échelle temporelle relativement courte. Cette situation est d'autant plus préoccupante pour les milieux arides dédiés au pastoralisme. Un surpâturage des terres de parcours combiné à des précipitations faibles et erratiques menacent la pérennité de la végétation et la sécurité alimentaire des populations locales qui dépendent de ces milieux pour leur survie. Pour ces raisons, il est primordial d'acquérir des connaissances sur le fonctionnement écologique des milieux arides et de s'attarder à leur devenir. Au cours de mon projet de doctorat, je me suis penché sur les relations entre les conditions environnementales et l'écologie fonctionnelle des communautés de plantes et de fourmis des steppes de l'Oriental marocain. Dans un premier temps, je me suis intéressé au changement dans la composition fonctionnelle de la végétation en fonction de l'aridité et du pâturage. Une approche fonctionnelle permet de suggérer des mécanismes explicites de l'assemblage des communautés en plus de pouvoir être utilisée peu importe les espèces et le type de milieu considérés. Les résultats ont mis en évidence trois groupes de traits fonctionnels qui rappellent la théorie C-S-R de Grime. La végétation a montré une stratégie d'évitement du pâturage plutôt qu'une réponse de résistance au pâturage en favorisant les espèces annuelles à croissance rapide, de petite taille et avec des graines de faible masse. L'aridité a sélectionné une végétation composée d'espèces ligneuses avec des feuilles succulentes et une teneur élevée en carbone isotopique 13. Malgré l'existence de corrélations significatives entre les traits fonctionnels et l'environnement, la force de ces corrélations était relativement faible. Le gradient d'aridité et le temps de mise en défens des exclos considérés n'étaient probablement pas suffisamment contrastés afin d'observer un filtrage environnemental net. Dans un deuxième temps, j'ai utilisé un modèle statistique novateur (MaxEnt) pour prédire les abondances relatives des espèces des communautés de plantes actuelles dans un contexte de changements climatiques. On prévoit une augmentation de l'aridité dans l'Oriental marocain pour la période 2080-2099. Le modèle MaxEnt s'appuie sur un principe clé de l'écologie fonctionnelle, soit le filtrage environnemental par les traits, et sur des outils empruntés à la mécanique statistique et à la théorie de l'information pour faire de telles prédictions. Les résultats ont montré que les prédictions d'abondances relatives des communautés actuelles sont plus précises lorsque les traits agrégés observés sont utilisés. Pour la période 2080-2099, le modèle MaxEnt suggère que les communautés passeront d'une dominance d'espèces annuelles à des espèces ligneuses tolérantes au stress. Ceci suggère un processus de désertification. Les prochaines études devront se pencher sur la définition du pool d'espèces et sur l'évolution des activités pastorales. Finalement, j'ai intégré les traits fonctionnels de fourmis à l'étude des communautés de plantes afin de suggérer une approche multitrophique à l'assemblage des communautés. Les résultats ont montré que la végétation a un effet direct sur la structure des communautés de fourmis et que l'environnement a un effet indirect sur ces mêmes communautés via la végétation. Ce modèle était supporté autant par les données de composition taxonomique que par les données de composition fonctionnelle des plantes et des fourmis. Ceci constitue un développement pour l'écologie des communautés, car ce type d'approche fonctionnelle est encore peu utilisé en écologie animale. Le développement d'un cadre conceptuel similaire à celui de la théorie C-S-R de Grime pourrait être très utile pour mieux comprendre les mécanismes d'assemblage des communautés animales.

The literature, long-term temporal data sets, and spatial data points surrounding livestock watering pints within rangeland communities wearer used to examine common assumptions and concepts used in models of vegetation dynamics. Of the stability concepts examined, the constancy concept was invalidated using long-term relict area data, whereas long-term data following disturbance indicated that sagebrush-dominated areas could be considered as resilient under circumstances examined. Pioshpere as well as relict and post-disturbance data indicated that the individualistic notion of vegetation change was favored. Species showing the most predictability (based on repeatability of abundance curves, and spatial and temporal serial correlations) were sagebrush and cheatgrass, specifically in their response to fire. Bunchgrasses were characterized by their persistence in abundance under all circumstances (grazing disturbance and fire disturbance) except long-term absence of fire. This, together with the support of the individualistic notion of plant community change, suggests examining change on a vital attributes basis with kinetic framework. It was found that most models of vegetation dynamics could be validated, because of the multivariate nature of the data examined. A closer examination of the requirements of range management for a models to be used as a basis for range condition and trend analysis determined that three basic model types are needed by range managers: First, a conceptual model for marrying divergent perspectives of different rangeland user groups; send, mechanistic models for the exploration of vegetation dynamics; and third, expert systems for information transfer. Further emphasis was placed on deriving a suitable conceptual models subscribing to the Objective of linking vegetation dynamics, range management, and monitoring within sagebrush-dominated areas. Important characteristics (derived from the data sets examined) included in this model are the recognition of static and dynamic plant communities, swathes of change rather than predefined pathways, the individualistic nature of change, and the recognition of spatial heterogeneity. Predictability at various scales was incorporated by focusing on cheatgrass and sagebrush, since these species indicated the most repeatable dynamics observed in the data sets, and were “common denominator”’ species (thus overcoming the problem of spatial heterogeneity). The undesired effects of cheatgrass and sage brush on other growth-forms suggest that monitoring and managing these undesired species may be a proactive way of managing for a desired plant community incorporating a balance of growth-forms.

The Eurasian steppe belt is the vastest grassland region worldwide, stretching approximately 8000 km from the Pannonian Basin in the west to the Amur river in the east, altogether covering more than 8 million km2. Due to its size and location, the vegetation of this biome was under strong influence of past climatic fluctuations that reached their peak in the Pleistocene. Fossil record of different steppe-associated flora and fauna places the onset of Eurasian open grasslands into Central Asia from which the grasslands spread westwards around 20 MYA and reached the East European Plain first in the Late Miocene around 8 MYA. However, while useful as a proxy to infer past vegetation patterns, fossils suffer from low-resolution power and thus fail to elucidate more detailed picture of the onset and the development of the Eurasian steppe belt flora. The working hypothesis driving the whole project was that molecular signals in typical steppe plant species reflect the climate-landscape history of the steppe and the biogeographic dynamics of steppe taxa and thus allow for a much finer resolution of the history of the steppe belt in comparison to floristic and fossil-based methods. By applying a plethora of different phylogenetic, phylogeographic and biogeographic methods, I first investigated the evolutionary history of four unrelated Brassicaceae taxa that can be nowadays found in the Eurasian steppe belt. Second, I tried to draw parallels with the climate-landscape history of the Eurasian steppe belt as inferred from the fossil record to test the above-mentioned working hypothesis. The first case study dealt with the economically important Brassicaceae genus Camelina, with an emphasis on C. microcarpa, which can be found across the whole Eurasian steppe belt. I uncovered that this taxon’s contemporary uninterrupted distribution was split along the north coast of the Caspian Sea approximately 1 MYA, dating back to the Apsheron and Baku transgression events. During this time period, a polyploidisation event took place giving rise to a new hexaploid taxon and subsequently preventing gene flow after the regression of the Caspian Sea. The second case study investigated the evolutionary history of Schivereckia, which exhibits a highly disjunct distribution along the East European Plain and the Balkans and the whole Schivereckia clade that can be nowadays found either at higher latitudes in the subarctic zone or mountain ranges of the northern hemisphere. My analyses placed the radiation of this clade at the beginning of Pleistocene, when low temperatures promoted speciation and radiation of cold-adapted flora and fauna. The study showed that the highly disjunct distribution of Schivereckia podolica mirrors the Pleistocene refugial areas of different ages and points towards a close evolutionary relationship between contemporary steppe and tundra flora. A third case study focussed on Sisymbrium. I uncovered that while Sisymbrium did not evolve in the Eurasian steppe belt, it invaded this area as well as the Mediterranean several times independently throughout the Pleistocene in the south-north and east-west trajectory, respectively. This study proved yet again how variable the Eurasian steppe flora is in terms of evolutionary onset and that many of steppe elements originated in its adjacent areas in the south. These then subsequently migrated to the Eurasian steppe belt only after it was already fully developed either towards the west into the Euro-Siberian steppe or to the east into the Mongol-Chinese steppe. The fourth case study investigated the evolutionary history of Capsella. A GBS-based approach was proven successful for inferring biological history of young taxa. I showed that contemporary steppe element Capsella orientalis invaded the Eurasian steppe belt long after it was already developed in the south-north trajectory. I acquired further insights in the evolutionary history of its cosmopolitan sister tetraploid C. bursa-pastoris and placed the origin of the whole genus into the late Pliocene continental Asia. All studies showed the importance of a taxon sample and offered an alternative nesting dating approach for reliable calibration in cases where no fossil record could be obtained. In conclusion, evolutionary history of all four investigated taxa were shaped by the same environmental changes that played a major role in the biological history of the Eurasian steppe belt. Overall, our working hypothesis that molecular signals in typical steppe plant species reflect the climate-landscape history of the steppe and the biogeographic dynamics of steppe taxa, turned out to be correct.

No description available.

Partant de l’exemple des steppes sud-oranaises, nous avons essayé de démontrer l’existence d’un modèle nomade de la pratique spatiale ainsi que son évolution à travers le temps, avec ses conséquences visibles et substantielles dans cette portion de l’espace algérien. Car les relations que les nomades entretiennent avec leur espace s’établissent selon un système de significations propre à cette société, dont il traduit l’histoire, le parcours, les expériences, les valeurs et les imaginations… au total une culture qui lui est propre et qui, en interférant avec les caractéristiques et les atouts du milieu naturel, donne naissance à un paysage. Or, la société nomade ainsi que son mode d’utilisation et d’adaptation à l’espace fait l’objet depuis une trentaine d’années d’une restructuration qui a provoqué des mutations du genre de vie et une recomposition des territoires ancestraux. La sédentarisation demeure un élément essentiel de ce changement qui a ses répercussions sur l’espace en général et sur celui des sédentaires en particulier. Quoiqu’il perdure, le nomadisme se transforme, prenant d’autres formes et utilisant d’autres outils. Peut-on parler d’une adaptation ou d’une résidualité conduisant à la décadence et finissant par une disparition ? L’espace perçu, vécu et représenté des nomades est-il pris en compte dans la gestion et l’aménagement de l’environnement/territoire ? Ce qui est identifié pour les groupes nomades de la steppe sud-oranaise peut servir d’illustration pour évoquer certains processus communs à plusieurs cultures, ou de point de comparaison avec d’autres sociétés, notamment de chasseurs-cueilleurs, autres nomades confrontés à une culture et un modèle allogènes et puissants. / From the example of the steppes of South Oran, we tried to demonstrate a model of nomadic space practice and its evolution through time, with its own visible and substantial consequences in a portion of Algerian’s space. The relation between nomads and their space settled in a specific meaning system. Its history, career, experiences, values and imaginations… in short, a culture of its own is arise from that relation, which by interfering with the features and advantages of the natural environment rise to a landscape. However, the nomadic society and its mode of space’s use and adaptation have been restructuring since the last thirty years. This restructuration has caused several mutations in their way of life and led to the reconstruction of ancestral territories. The settlement remains an essential element of those changes which impact on space. Although nomadism continues, it turns into other form and other tools. Can we speak of an adaptation or a residuality leading to decadence and ending with a loss? The Nomad’s perceived lived and represented space, is it taken into account in the management and development environment/territory? This identified, the south Oran nomadic society can serve as an illustration to evoke some process common to many cultures, or to compare it with other societies, such as hunter-gathers, others nomads facing an alien culture and powerful models.

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 270 / Mit 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 270

Le Moustérien se présente au Proche-Orient comme un technocomplexe qui associe des tendances générales dans la succession des industries lithiques, à une diversité technique difficile à cerner dans un cadre unitaire. Dans cette aire du bassin méditerranéen, témoin de dynamiques de peuplement complexes entre les continents, les modèles de la variabilité du Moustérien ont été élaborés à partir des sites de la zone côtière. Les sites de la zone steppique ont toujours été interprétés en relation à ces modèles. Pourtant, les données semblent témoigner d’une diversité technique plus importante. Nous proposons une lecture géographique de cette diversité à partir de la comparaison de trois séquences : Umm el Tlel, dans l’aire semi-aride de Syrie ; Yabroud I, dans les montagnes au nord de Damas ; et Ksar ‘Akil, sur la côte libanaise. A l’intérieur de la mosaïque paléoenvironnementale qui caractérise le Proche-Orient, chacun de ces sites rend compte d’un milieu particulier et permet d’appréhender la question des changements techniques dans différents contextes. L’analyse technologique met en évidence que, au-delà de la continuité d’un système de débitage unique, le système Levallois, chaque industrie témoigne d’objectifs et de méthodes différentes. La comparaison montre que les changements techniques ne se produisent pas partout de la même façon : à Umm el Tlel et Yabroud I, chaque couche présente une industrie différente, alors qu´à Ksar ‘Akil, une continuité technique est partagée par les industries de plusieurs couches, les changements se produisant à travers de grands ensembles successifs dans le temps. Nous proposons que ces différentes modalités de changements dans le temps soient liées à une diversité techno-culturelle plus importante des populations qui se sont succédées dans la zone steppique que dans la zone côtière. Des dynamiques d’occupation spécifiques de l’espace macro-régional sont alors proposées, qui peuvent expliquer ces différences. / The Near Easter Mousterian technocomplex shows some large chronological trends in the lithic industry changes, and also an internal technical diversity which is difficult to explain in a homogeneous framework. In this area of the Mediterranean basin, place of complex population dispersals between the continents, the first definitions and models of Mousterian variability, have been developed starting from sites in the coastal area. The steppic semi-arid area has always been interpreted using this model framework, but the data we have in this area suggest a deeper technical diversity. We introduce a geographical perspective to go through this technical diversity, based on the comparison of three Mousterian sequences: Umm el Tlel, in the semi-arid area of Syria, Yabroud I, in the mountainous area in the north of Damascus, and Ksar ‘Akil, on the Lebanese coast. In the paleoenvironmental mosaic which is the peculiarity of the Near Eastern region, each one of these sites belongs to a specific milieu, and documents the issue of technological changes in different contexts. Technical analyses show the continuity of the same production system, the Levallois débitage, shared by the industries of the three sites. In a comparative perspective, technical changes don’t appear at the same rhythm in the three sequences: in Umm el Tlel and Yabroud I, each layer has a different technological organisation; on the other side at Ksar ‘Akil a technical continuity is shared by several layers and changes appear following progressive trends during time. We suggest that these different change modalities are the result of a deeper techno-cultural diversity of human groups populating the steppic area. Specific settlement dynamics of the macro-regional space are suggested to explain these differences.

Habitat structure is an important driver of many ecological patterns and processes, but few studies investigate whether habitat structure interacts with other environmental variables to affect community dynamics. The main objective of this study was to disentangle the relative importance of prey availability and shrub architecture on the distribution, abundance, and biodiversity of spiders of northern Utah, USA. We conducted field experiments which focused on: (1) describing the importance of these factors on spider community organization, (2) specifically evaluating whether prey availability mediates the relationship between shrub architecture and spider abundance and biodiversity, and (3) investigating spider and prey responses to manipulations of surrounding vegetation structures. For the first two experiments, big sagebrush shrubs were randomly assigned to six experimental treatments: two levels of prey attractant (shrubs were either baited or not baited) and three levels of foliage density (low, natural/control, or high). The purpose of manipulating both prey availability and shrub architecture was to delineate their significance to spiders. For the last experiment, changes in these factors were investigated at two different levels of spatial context (a single manipulated shrub surrounded by untreated shrubs vs. a manipulated shrub surrounded by a patch of similarly treated shrubs). We found both prey availability and shrub architecture directly influenced patterns of spider abundance and species richness and that spider species diversity and community composition varied in response to shrub architecture alone. Preferences of some spiders for certain shrub types likely reflect differences in foraging strategies or the substrate required to support different types of webs. We also demonstrate that spider response to shrub architecture is the result of multiple processes (i.e., a combination of direct and indirect effects via prey availability) and that surrounding vegetation structures affect spider abundances on shrubs. In addition, prey composition varied among different shrub foliage density treatments, but only when surrounding vegetation structures were also manipulated. More generally, this study suggests that ecological responses to habitat structure are in part mediated by associated variables and the significance of shrub architecture varies depending on the organisms examined and the spatial scale to which they respond most strongly.