Dynamics in Organic Matter Processing, Ecosystem Metabolism and Trophic Sources for Consumers in the Mara River, Kenya Frank Onderi Masese DYNAMICS IN ORGANIC MATTER PROCESSING, ECOSYSTEM METABOLISM AND TROPHIC SOURCES FOR CONSUMERS IN THE MARA RIVER, KENYA Frank Onderi Masese Thesis committee Promotor Prof. Dr Kenneth A. Irvine Professor of Aquatic Ecology and Water Quality Management UNESCO-IHE Institute for Water Education Delft, The Netherlands Co-promotors Prof. Dr Michael E. McClain Professor of Ecohydrology UNESCO-IHE Institute for Water Education Delft, The Netherlands Dr Gretchen M. Gettel Senior Lecturer in Aquatic Biogeochemistry UNESCO-IHE Institute for Water Education Delft, The Netherlands Other members Prof. Dr Wolf Mooij, Wageningen University Prof. Dr Robert Naiman, University of Washington, USA Prof. Dr Steve Ormerod, Cardiff University, UK Dr Martin Pusch, Leibniz-Institute of Freshwater Ecology and Inland Fisheries, Germany This research was conducted under the auspices of the SENSE Research School for Socio-Economic and Natural Sciences of the Environment DYNAMICS IN ORGANIC MATTER PROCESSING, ECOSYSTEM METABOLISM AND TROPHIC SOURCES FOR CONSUMERS IN THE MARA RIVER, KENYA Thesis submitted in fulfilment of the requirements of the Academic Board of Wageningen University and the Academic Board of the UNESCO-IHE Institute for Water Education for the degree of doctor to be defended in public on Wednesday, 11 March 2015 at 3 p.m. in Delft, the Netherlands by FRANK ONDERI MASESE Born in Kisii, Kenya CRC Press/Balkema is an imprint of the Taylor & Francis Group, an informa business © 2015, Frank Onderi Masese All rights reserved. No part of this publication or the information contained herein may be reproduced, stored in a retrieval system, or transmitted in any form or by any means, electronic, mechanical, by photocopying, recording or otherwise, without written prior permission from the publishers. Although all care is taken to ensure the integrity and quality of this publication and information herein, no responsibility is assumed by the publishers or the author for any damage to property or persons as a result of the operation or use of this publication and or the information contained herein. Published by: CRC Press/Balkema PO Box 11320, 2301 EH Leiden, The Netherlands e-mail: [email protected] www.crcpress.com – www.taylorandfrancis.com ISBN 978-1-138-02815-9 (Taylor & Francis Group) ISBN 978-94-6257-209-6 (Wageningen University) Dedication To the loving memory of my mother, the late Miriam Gesare Masese v analysis. Thank you Amanda for the many discussions we had by the river and the Acknowledgements encouraging words. I appreciate Dr Fidelis Kilonzo (Kenyatta University, Kenya and Leaping across streams every week-day for school nearly three decades ago, my early days in UNESCO-IHE), Dr Paolo Paron and Veronica Minaya (UNESCO-IHE) for their assistance formal education were intimate with nature. In these streams, I and my childhood friends with the maps used in this dissertation. I would like to express my sincere gratitude to Dr Erik fished with baskets and hook and line, and I was always intrigued by the diversity of aquatic de Ruyter van Steveninck for the Dutch translation of the summary of this dissertation. macrofauna. Even though arduous and winding, my academic journey has gone full circle and taken me back to streams. The success has however been achieved with a dose of I appreciate colleagues at UNESCO-IHE such as Jeremiah Kiptala, Dr Chol Deng Thon Abel, appreciation, and I thank God for His providence and the many people to whom I owe so Veronica Minaya, Graciela Alvarez, Adeboye Omatayo, Amadou Keita, Dr George Lutterodt, much. Dr Fidelis Kilonzo and Dr Njenga Mburu with whom I shared frustrations, excitements, sought informal advice and relaxed with away from the rigors of dissertation work; they made This dissertation has been made possible by the guidance and support of my promoters, the PhD experience to be social, enjoyable and fulfilling. I am grateful to the many graduate mentors, colleagues, financial assistance from the Dutch Ministry of Foreign Affairs and students whom I worked with in the Mara and the enriching discussions we had; Elvirah support and encouragement from my wife. Firstly, I would like to thank my promoter and Riungu and Zipporah Gichana (University of Eldoret), Evance Mbao, Cheplabatt Kabon and mentor Prof. Michael E. McClain (UNESCO-IHE, The Netherlands), for his support, Audrey Tsitsiche (Egerton University), and Jessica Salcedo Borda and Anne Lilande mentoring and advice throughout the entire PhD program, as well as for giving me the (UNESCO-IHE). I am also grateful for the dedicated and timely assistance of Ms Jolanda opportunity to work on this project. I would also like to thank Michael for always being Boots at the UNESCO-IHE fellowship office. approachable and happy to discuss ideas and challenges, and for providing timely and constructive reviews of manuscripts and draft chapters. I would like to thank my promoter I appreciate the funding sources that made this dissertation possible. Firstly, the Dutch Prof. Kenneth Irvine and supervisor Dr Gretchen Gettel (UNESCO-IHE, The Netherlands), Ministry of Foreign Affairs through UNESCO-IHE Partnership Research Fund (UPaRF) for for their helpful advice, comments and reviews of manuscripts and dissertation chapters and funding the MaraFlows project through which I was offered this PhD fellowship. I would like the wealth of experience and different approaches they brought to my advisory board. I to appreciate the USAID MSc scholarships through the GLOWS project that contributed to appreciate the guidance and support offered by Dr Nzula Kitaka (Associate Professor) and Dr some MSc studies under my dissertation work, and the partial support provided by AFRIVAL Julius Kipkemboi (Associate Professor) of Egerton University, Kenya. I also appreciate the (ERC-Stg 240002) for the stable isotopes analysis at KU Leuven. I would also like to input offered by Prof. Steven Bouillon (KU Leuven, Belgium) and Dr. Katya Abrantes (James acknowledge the Kenya National Council for Science and Technology for granting me and Cook University, Australia) for their advice and assistance during analysis of stable isotopes my supervisory team permission to conduct this research. I appreciate the TransMara data. Conservancy and the Narok County Council (Narok County) for granting me access into the Mara Triangle and Masai Mara National Reserve, respectively. I acknowledge the assistance offered by many people during field work and in the laboratory which made the compilation of this dissertation a success. To Kimutai Kitur and Robertson To my lovely wife Nyatichi, thank you so much for all your support and patience, and for Marindany, thanks for spending days in the field collecting water quality data and samples, reminding me that there was more to life than just this dissertation. Special and profound fish, bugs, and plants from the crocodile and hippo infested Mara River. Thanks to Dr Phillip appreciation for taking care of our son Bartundo during the long days spent away from home. O. Raburu (Associate Professor, University of Eldoret) for the support, guidance and Special thanks to my father, brothers and sisters for their love and encouragement. logistical assistance during field work. I acknowledge Dr William Ojwang and Chrisphine Nyamweya (KMFRI, Kisumu) and their team for assistance during fish sampling and gut content analysis. Special thanks to Lubanga Lunaligo and David Namwaya (University of Delft, November 2014 Eldoret) for assistance during lab work; I cannot forget to mention the commitment with Frank Onderi Masese which they helped open guts of bugs and sort litter. I thank Zita Kelemen (KU Leuven), Amanda Subalusky and Dr Glendon Hunsinger (Yale University, CT, USA) for stable isotope vi vii analysis. Thank you Amanda for the many discussions we had by the river and the encouraging words. I appreciate Dr Fidelis Kilonzo (Kenyatta University, Kenya and UNESCO-IHE), Dr Paolo Paron and Veronica Minaya (UNESCO-IHE) for their assistance with the maps used in this dissertation. I would like to express my sincere gratitude to Dr Erik de Ruyter van Steveninck for the Dutch translation of the summary of this dissertation. I appreciate colleagues at UNESCO-IHE such as Jeremiah Kiptala, Dr Chol Deng Thon Abel, Veronica Minaya, Graciela Alvarez, Adeboye Omatayo, Amadou Keita, Dr George Lutterodt, Dr Fidelis Kilonzo and Dr Njenga Mburu with whom I shared frustrations, excitements, sought informal advice and relaxed with away from the rigors of dissertation work; they made the PhD experience to be social, enjoyable and fulfilling. I am grateful to the many graduate students whom I worked with in the Mara and the enriching discussions we had; Elvirah Riungu and Zipporah Gichana (University of Eldoret), Evance Mbao, Cheplabatt Kabon and Audrey Tsitsiche (Egerton University), and Jessica Salcedo Borda and Anne Lilande (UNESCO-IHE). I am also grateful for the dedicated and timely assistance of Ms Jolanda Boots at the UNESCO-IHE fellowship office. I appreciate the funding sources that made this dissertation possible. Firstly, the Dutch Ministry of Foreign Affairs through UNESCO-IHE Partnership Research Fund (UPaRF) for funding the MaraFlows project through which I was offered this PhD fellowship. I would like to appreciate the USAID MSc scholarships through the GLOWS project that contributed to some MSc studies under my dissertation work, and the partial support provided by AFRIVAL (ERC-Stg 240002) for the stable isotopes analysis at KU Leuven. I would also like to acknowledge the Kenya National Council for Science and Technology for granting me and my supervisory team permission to conduct this research. I appreciate the TransMara Conservancy and the Narok County Council (Narok County) for granting me access into the Mara Triangle and Masai Mara National Reserve, respectively. To my lovely wife Nyatichi, thank you so much for all your support and patience, and for reminding me that there was more to life than just this dissertation.