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The Return of the Mammoth Steppe? – Rewilding in Yakutia and the Actual Impact of Large Herbivore Grazing on Vegetation

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The Return of the Mammoth Steppe? – Rewilding in Yakutia and the Actual Impact of Large Herbivore Grazing on Vegetation DISSERTATION The return of the mammoth steppe? – Rewilding in Yakutia and the actual impact of large herbivore grazing on vegetation By Jennifer Reinecke Dresden, 2019 The return of the mammoth steppe? – Rewilding in northeastern Yakutia and the actual impact of large herbivore grazing on vegetation Dissertation zur Erlangung des akademischen Grades Doctor rerum naturalium (Dr. rer. nat.) vorgelegt dem Bereich Mathematik und Naturwissenschaften der Technischen Universität Dresden von Jennifer Reinecke (M.Sc.) geboren am 18.12.1983 in Berlin, Deutschland Eingereicht am 24.05.2019 Verteidigt am 15.11.2019 Gutachter: Prof. Dr. Karsten Wesche Prof. Dr. Christine Römermann Prüfungskommission: Prof. Dr. Christoph Neinhuis Prof. Dr. Christine Römermann Prof. Dr. Goddert von Oheimb Prof. Dr. Karsten Wesche ii Die Dissertation wurde in der Zeit von Juni 2014 bis Mai 2019 unter Betreuung durch Prof. Dr. Wesche im Senckenberg Museum für Naturkunde, Görlitz, angefertigt. iii iv Eidesstattliche Erklärung Hiermit versichere ich, Jennifer Reinecke, dass ich die vorliegende Arbeit mit dem Titel ÄThe return of the mammoth steppe? ± rewilding in Yakutia and the actual impact of large herbivore grazing on vegetation ³ ohne unzulässige Hilfe Dritter und ohne Benutzung anderer als der angegebenen Hilfsmittel angefertigt habe. Die aus fremden Quellen direkt oder indirekt übernommenen Gedanken sind als solche kenntlich gemacht. Die Arbeit wurde bisher weder im Inland noch im Ausland in gleicher oder ähnlicher Form einer anderen Prüfungsbehörde vorgelegt. Ich erkenne die Promotionsordnung der Fakultät Mathematik und Naturwissenschaften der Technischen Universität Dresden vom 23.02.2011 an. Dresden, 15.05.2019 Jennifer Reinecke ǀ Vorwort 1DFK $OH[DQGHU YRQ +XPEROGW 9RUUHGH ]XU $XVJDEH YRQ Ä$QVLFKWHQ GHU 1DWXU³ Ä6FKFKWHUQEHUJHEHLFKGHP3XEOLNXPHLQH5HLKHYRQ$UEHLWHQGLHLP$QJHVLFKW JURHU 1DWXUJHJHQVWlQGH >«@ in den Steppen von [Sibirien] entstanden sind. Einzelne Fragmente wurden an Ort und Stelle niedergeschrieben und nochmals nur in ein Ganzes zusammengeschmolzen. Überblick der Natur im großen, Beweis von dem Zusammenwirken der Kräfte, Erneuerung des Genusses, welchen die unmittelbare Ansicht der [Wildnis] dem fühlenden Menschen gewährt, sind die Zwecke, nach denen ich strebe. Jeder Aufsatz sollte ein in sich geschlossenes Ganzes ausmachen, in allen sollte eine und dieselbe Tendenz sich gleichmäßig ausVSUHFKHQ>«@5HLFKWXPGHU1DWXU veranlasst Anhäufung einzelner Bilder, und Anhäufung stört die Ruhe und den 7RWDOHLQGUXFNGHV*HPlOGHV>«@0|JHQPHLQH$QVLFKWHQGHU1DWXUWURW]GLHVHU)HKOHU welche ich selber leichter rügen als verbessern kann, dem Leser doch einen Teil des Genusses gewähren, welchen ein empfänglicher Sinn in der unmittelbaren Anschauung ILQGHW³ ǀŝ Für meine kleine Nomadenschwester ǀŝŝ Contents I. Acknowledgements xi Chapter 1: General Introduction 1 1.1. Large mammal extinctions and the onset of the Anthropocene 1 1.2. 7KH´PDPPRWKVWHSSH´ 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 7 of the mammoth steppe 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 ǀŝŝŝ Chapter 3: Extrazonal steppes and other temperate grasslands of northern Siberia í3K\WRVRFLRORJLFDOFODVVLILFDWLRQDQGHFRORJLFDOFKDUDFWHUL]DWLRQ 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 &KDSWHU:RRGODQGVDQGVWHSSHV3OHLVWRFHQHYHJHWDWLRQLQ<DNXWLD¶V most continental part recorded in the Batagay permafrost 82 sequence 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 113 and environment throughout the sequence 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: 124 Pleistocene survivors or Holocene immigrants? 4.4.3. Climatic implications 126 4.5. Conclusions 129 ŝdž 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 157 - and their proposed role in the mammoth steppe 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 dž I. Acknowledgements The project ³7KH HQG RI WKH 3OHLVWRFHQH WXQGUD VWHSSH ± interactions between vegetation, climate and large herbivores in Beringia during the late Quaternary (TUNDRA-67(33( ´ and my three-year PhD-position were funded by the DFG (KI 849/4-1, WE 2601/9-1). I want to thank Frank Kienast for coming up with this very interesting research idea and for arranging and proposing the project to the DFG together with Karsten Wesche. I am truely grateful to both of them for giving me this challenging yet exciting opportunity. I can also honestly and very gladly say that I found a very dedicated mentor in Karsten Wesche. We had many interesting scientific discussions and inspiring talks about the scientific world and beyond, despite the little time this same scientific system leaves supervisors to dedicate to their students. I am also very thankful to the co-authors, reviewers and editors of my papers for for valuable questions and comments, which not only improved my manuscripts, but my scientific understanding. Kseniia Ashastina and Elena Troeva are my heros of our expeditions! I probably only survived the wild Siberian outdoors and hardy Russian people because of their professional organization and communication skills. Elena and her family gave us a very warm welcome to Yakutia and I owe it to her networking and organizational efforts that we were able to properly collect our data and savely bring it home to Germany. Our second expedition would not have been possible without the invitation and extensive logistic help of the Zimov family in the Northeast Science Station. I thank Nikita Zimov for the welcoming support of our research at their incredible wildlife park, the Pleistocene Park. I am grateful for controversial discussions on rewilding with Sergey Zimov and Sergey Davidov, which made me dig deeper into the different arguments. And I thank all the workers of Pleistocene Park for keeping us save all day from the bison in the field and taking us to all the impossible sites we wanted to visit. I also thank the Yakutian government and the workers of the Bisonary for their support regarding accommodation, information and supervision in their wildlife park. I even want to thank the taxi driver who took us to the steppes around Yakutsk for his help with cutting biomass out of boredom and for incredulously asking the right question to keep my eyes IL[HGRQVFLHQFHLQWKHKDUGVKLSWKDWILHOGZRUNFDQEH ³:K\GRQ¶W\RXMXVWFXWDQ\JUDVV VDYHWLPHDQGJRKDYHDEHHUIRUWKHUHVWRIWKHGD\"´ The large amounts of samples, which I brought with me from Siberia, would have been impossible to handle without the dedicated work of many collegues. In the laboratories of Senckenberg Natural History Musuem Görlitz I especially want to thank Michaela Schwager and Sebastian Moll, as well as Carmen Trog for their extensive logistic help. I am grateful to Volker Otte and his co-workers, who helped to collect lichen and mosses in the first expedition and identified them back home.
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