Zurich Open Repository and Archive University of Zurich Main Library Strickhofstrasse 39 CH-8057 Zurich www.zora.uzh.ch Year: 2015 Reconstructing the potential natural vegetation and soils of the high Andes Sylvester, Steven Paul Posted at the Zurich Open Repository and Archive, University of Zurich ZORA URL: https://doi.org/10.5167/uzh-119640 Dissertation Published Version Originally published at: Sylvester, Steven Paul. Reconstructing the potential natural vegetation and soils of the high Andes. 2015, University of Zurich, Faculty of Science. Reconstructing the potential natural vegetation and soils of the high Andes Dissertation zur Erlangung der naturwissenschaftlichen Doktorwürde (Dr.sc.nat.) vorgelegt der Mathematisch-naturwissenschaftlichen Fakultät der Universität Zürich von Steven Paul Sylvester aus England Promotionskomitee: PD. Dr. Michael Kessler (Vorsitz) Prof. Dr. Peter Linder Dr. Colin Hughes Zürich, 2015 Contents ACKNOWLEDGMENTS ............................................................................................................................. 5 ZUSAMMENFASSUNG ............................................................................................................................. 7 SYNOPSIS ................................................................................................................................................. 9 PROLOGUE ............................................................................................................................................ 11 THE QUESTION: ARE THE HIGH-ELEVATION ECOSYSTEMS OF THE ANDES NATURAL? ........................ 11 THE TASK: OBTAINING BASELINE DATA ................................................................................................ 14 THE SOLUTION: INACCESSIBLE LEDGES................................................................................................. 15 THE FINDINGS ....................................................................................................................................... 16 CHAPTER 1: Inaccessible ledges as refuges for the natural vegetation of the high Andes. ............. 31 CHAPTER 2: Inaccessible Andean sites reveal human induced weathering in grazed soils. ............. 45 CHAPTER 3: Shifting perspectives on naturalness in mountain ecosystems. ................................... 73 CHAPTER 4: Elevational patterns of Polylepis tree height (Rosaceae) in the high Andes of Peru: role of human impact and climatic conditions ..................................................................... 139 CHAPTER 5: The World's highest vascular epiphytes found in the Peruvian Andes. ........................ 153 CHAPTER 6: An updated checklist and key to the open-panicled Poa’s (Poaceae) of Peru including 3 new species, Poa ramoniana, Poa tayacajaensis and Poa urubambensis. ................ 163 CHAPTER 7: Bartsia lydiae, a new species of Bartsia sect. Laxae (Orobanchaceae) from the southern Peruvian Andes with a revised key to Bartsia sect. Laxae. ........................... 193 CHAPTER 8: A new species of Moranopteris (Polypodiaceae) from inaccessible ledges in the high Andes of Peru. ............................................................................................................... 201 CHAPTER 9: Gentianella viridiflora (Gentianaceae), a new green-flowered Gentianella species from the Peruvian Andes. ...................................................................................................... 213 CHAPTER 10: The World’s smallest Campanulaceae: Lysipomia mitsii sp.nov. .................................. 221 OUTLOOK AND FUTURE RESEARCH ...................................................................................................... 241 CURRICULUM VITAE .............................................................................................................................. 245 3 4 ACKNOWLEDGEMENTS First and foremost, God deserves all the glory for this. Thanks to this PhD, I met my soulmate and wife, Mitsy, the best climbing partner, friend and accomplice that one could ever ask for, who’s patience, wisdom, stamina and general awesomeness I have not met the like of in any other person that I know. This project would not have been in any way as successful without Mitsy’s help and she deserves a large part of the credit. I would like to express my deepest thanks to Michael Kessler, my fantastic PhD supervisor, who had the visionary idea behind this project, whilst I had only voiced my eagerness to combine botany with climbing in South America. I thank Michael for having the belief in me that I could perform this research and for teaching me a lot and putting up with my escapades over the last 5 years. The Leverhulme Trust deserves an immense thanks for their generous funding which allowed the majority of field research to be done. This should not be understated, as most other funders regarded the project to have a high chance of failure and were not prepared to take the risk. Thank you Leverhulme Trust for having the belief in me that I could perform this research. Following this, funding came more easily, and I am grateful to the Deutsche Forschungsgemeinschaft [grant numbers HE 6726 5/1; JU XX], and the Claraz-Schenkung fund for additional fieldwork funding which allowed me to extend my stay in Peru. The University of Zurich Forschungskredit, as well as additional funding from the budget of Michael Kessler and Elena Conti, are thanked for allowing me sufficient time in Zurich to publish the results of the fieldwork and to prepare this thesis. My brilliant colleagues, Felix Heitkamp and Hermann Jungkunst, helped me a lot with regards preparing chapters 1 to 3 and all soil aspects of the project. It’s amazing that Felix still speaks to me after my crazy driving on the Peruvian mountain roads and lowering him off cliffs. My new Peruvian family deserve a lot of thanks for their constant support. Many problems in Peru ranging from mind-numbing bureaucracy, logistical nightmares of fieldwork, and general problems whilst living in the country, were rapidly solved through the help of my father-in-law and uncle, Walter and Humberto Diaz Galdos, and my sisters-in- law, Patricia and Daniela Diaz Puente de la Vega. The unconditional support of my own family was felt through the entire project, and I really thank them for it. My parents, Paul and Barbara, and my grandmother, Jeanette, are especially thanked for all the help they gave. The LPB herbarium and staff were an immense help in giving me a workspace and providing all the relevant infrastructure so that I could spend many a merry month identifying specimens in La Paz. Following on from this, many specialists provided their knowledge and expertise. Juan Camilo Ospina, Simon Laegaard, Robbert Gradstein, Steve Churchill, Robert Soreng, Xenia Villavicencio, Michael Sundue, Michael Kessler, Colin Hughes, Ulf Molau, Simon Pfanzelt, Klaus Bernhard von Hagen, Fred Barrie, Zulma Rugulo and Hibert Huaylla are all thanked for help with specimen identification and for helpful discussion regarding species delimitation, with which I gained extra experience and knowledge of these interesting plant groups. Harrie Sipman and the Berlin Dahlem Botanical Garden and Botanical Museum are thanked profusely for allowing my wife, Mitsy, to stay at the Botanical Garden for extended periods to identify the lichen specimens and benefit from brilliant training in lichen taxonomy. Johanna Toivonen and Carlos Gonzales Inca provided 5 their skills in GIS and forest structure data from disturbed ecosystems and I am very grateful for their collaboration and hope it continues. Tatiana Erika Boza is greatly thanked for her help in measuring Polylepis trees whilst in Cusco as, without this help, some of the main discoveries of chapter 3 would not have been possible. ECOAN-Peru introduced us to some of the mountain communities and provided correspondence so that we were accepted by, at least some, of these high mountain people, whilst many others refused us entry or even tried to burn down our camps. Dr. Washington Galiano initially helped with specimen identification and was invaluable in his help obtaining the research and export permits. Giulia Attochi volunteered for a short time at the beginning of the project and is thanked for all her effort and sharing in the adventure. I would really like to thank Dr. Dirk Karger and his family for all their help and assistance with things ranging from helping us to settle in to Swiss life, offering us a place to stay and momentary relaxation through fondue and sledging. Hopefully we can repay this kindness in the future. Dr. Karger also was an invaluable help with statistics, managing R software, and discussion over general ecological concepts. I am very grateful to him for translating the synopsis into German. My great friends, Jakob and Heidi Frauenfelder, deserve a large amount of thanks for all the hospitality and help they have given during our stay in Zurich, and for showing us some really fantastic places. God bless you both. Jakob was also able to beautify the German translation of the synopsis. All the great people at the Institute of Systematic Botany are thanked for their friendship and help during the last few years. Particular mention goes to Colin Hughes and Peter Linder who agreed to be on my PhD committee and who provided help and advice on many issues. On my first trips to Zurich, Jurriaan M. de Vos, Danielle Cicuzza and Stefan Abrahamczyk are thanked for their great hospitality in