Recovery of a tropical rain forest over 30 years following silvicultural interventions Thesis submitted in partial fulfilment of the requirements of the degree Doctor rer. nat. of the Faculty of Environment and Natural Resources, Albert-Ludwigs-Universität Freiburg im Breisgau, Germany by Angela Luciana de Avila Freiburg im Breisgau, September 2016 Dean: Prof. Dr. Tim Freytag First examiner: Prof. Dr. Jürgen Bauhus Second examiner: Prof. Dr. Jaboury Ghazoul Third examiner: Prof. Dr. Carsten Dormann Date of thesis defence: 23rd January 2017 Statement of originality I hereby declare that this thesis has never been submitted to another examination commission in Germany or in another country for a degree in the same or similar form. This thesis contains no material previously published or written by another person except where due acknowledgement is made in the proper manner. Angela Luciana de Avila Freiburg, 29th September 2016 Statement of contributions and manuscripts This doctoral research was carried out between April 2012 and September 2016. I conducted most of the work myself, from the development of the research concept, data preparation and analyses to the writing of manuscripts, introduction and final chapters of this thesis. Three manuscripts were elaborated to be published in peer-reviewed scientific journals, as follows: de Avila, A.L., A.R. Ruschel, J.O.P. de Carvalho, L. Mazzei, J.N.M. Silva, J.do.C. Lopes, M.M. Araujo, C.F. Dormann, and J. Bauhus. 2015. Medium-term dynamics of tree species composition in response to silvicultural intervention intensities in a tropical rain forest. Biological Conservation, 191:577–586. de Avila, A.L., M. van der Sande, C.F. Dormann, L. Mazzei, M. Peña-Claros, L. Poorter, A.R. Ruschel, J.N.M. Silva, J.O.P. de Carvalho, and J. Bauhus. Biomass resilience to silvicultural interventions in a tropical rain forest: effects of disturbance intensity and tree community properties. Submitted for publication. de Avila, A. L., G. Schwartz, A.R. Ruschel, J.doC. Lopes, J.N.M. Silva, J.O.P. de Carvalho, C.F. Dormann, L. Mazzei, M. Soares, and J. Bauhus. 2017. Recruitment, growth and recovery of commercial tree species over 30 years following logging and thinning in a tropical rain forest. Forest Ecology and Management, 385: 225–235. Prof. Dr. Jürgen Bauhus supervised and participated actively in all stages of the work. Prof. Dr. Carsten Dormann helped with data analysis and development of manuscripts. Dr. José N. M. Silva, Dr. João O. P. de Carvalho and Dr. José. C. Lopes designed the experiment and accompanied it since 1981. In this thesis, they contributed to improve the manuscripts. Dr. Ademir R. Ruschel, Dr. Lucas Mazzei and Marcio Soares are responsible for the continued monitoring of the experiment and helped with data preparation and improvement of manuscripts. Prof. Dr. Maristela Araujo participated in the development of the first manuscript. Dr. Masha van der Sande coordinated the collection of functional trait data used in the third chapter and together with Prof. Dr. Marielos Peña-Claros and Prof. Dr. Lourens Poorter helped develop the concept and improve the second manuscript. Gustavo Schwartz contributed to develop the third manuscript. Data collection was carried out by the Forest Management Group of Embrapa and I participated in the last inventory in 2012. Lastly, five conference presentations were held and one additional publication resulted from a scientific collaboration, as follows: van der Sande, M., E.J. Arets, M. Peña-Claros, A.L. de Avila, A. Roopsind, L. Mazzei, N. Ascarrunz, B. Finegan, A. Alarcón, Y. Cáceres-Siani, J.C. Licona, A.R. Ruschel, M.Toledo, and L. Poorter 2016. Old-growth Neotropical forests are shifting in species and trait composition. Ecological Monographs, 86: 228-243. I dedicate this thesis to my parents José and Marli who taught me to love nature. Dedico esta tese aos meus pais José e Marli que me ensinaram a amar a natureza. Acknowledgements I thank God for giving me life and health that enable me to experience and learn from life at its fullest extent. Thanks for the greatness and beauty of the Amazonian or any other tropical rain forest which always inspired me to try to do something to help conserve these ecosystems. Doing this doctoral research was an enjoyable experience that made me grow both personally and professionally. This journey started in 2011, when after finishing my Master thesis on the regeneration of Araucaria forests in south-eastern Brazil, I wanted to do something concrete to help towards the conservation of Amazonian forests. As a forester, I believe in the potential of sustainably managing these ecosystems to combine the production of goods and the conservation of biodiversity and other associated benefits. As a scientist, I believe that science can and needs to contribute with generating knowledge to this purpose. I am grateful to Prof. Dr. Maristela Machado Araujo for her friendship and support. She helped me build the bridge towards my professional objective. I extend my gratitude to Prof. Dr. João Olegário Pereira de Carvalho who accepted my participation in such a long-term project, which provided the data to be evaluated in this thesis. I also thank Prof. Dr. Benno Pokorny who helped get in contact with my supervisor, Prof. Dr. Jürgen Bauhus, whom I am deeply grateful for believing in my doctoral proposal and on my potential to develop it. He instigated and inspired the development of new and interesting ideas and always motivated me to make the most out of this doctoral research, encouraging and supporting me throughout the whole process. I am also very grateful to my second supervisor, Prof. Dr. Carten Dormann, who taught me theories and tools for statistical analysis, instigated critical thinking and closely supported and motivated the development of this research. Thanks also to Prof. Dr. Jaboury Ghazoul for accepting to examine this doctoral thesis. I would like to extend my thanks to the many institutions that supported the development of this doctoral research. Embrapa Amazônia Oriental was the host institution of this project and I thank Embrapa for the possibility of working with such a valuable dataset. I thank the German Academic Exchange Service (DAAD) for financing my doctoral studies at the University of Freiburg (doctoral scholarship) and Mrs. Maria Salgado from DAAD office, who was always ready to help. I thank the Müller-Fahnenberg Foundation for supporting the field trip to accompany the inventory of permanent sample plots in 2012. I am thankful to the following institutions for supporting my participation in academic conferences and summer schools that surely helped in the development of this research and of my academic career: the Müller- Fahnenberg Foundation, the Sino-German Centre for Research Promotion, the Georg-Ludwig- Hartig foundation, the Council for Tropical and Subtropical Agricultural Research, the Wissenschaftliche Gesellschaft, the International Graduate Academy, and the graduate school “Environment, Society and Global Change”. Thanks also to the two last institutions for providing courses to improve professional skills. This work could not have been done without the active participation of many people. I am grateful to Dr. Ademir Roberto Ruschel for all the support provided throughout the development of this research. I also thank other members of the Embrapa Forest Management group that were always open and available to exchange ideas and help in the development of this research: Dr. Lucas Mazzei, Marcio Soares, Dr. Gustavo Schwartz, Dr. José do Carmo Lopes and Dr. José Natalino Macedo Silva. My gratitude extends to Lúcio Reginaldo Seixas and Nilson de Souza Carvalho that taught me a lot about the Amazonian forests during fieldwork and made that time fun. I thank Rubens and Vera for supporting me during my stops in Santarém for the travels between Belém and the study area, and Esther Muschelknautz for her support to me as a member of the Graduate School. I am thankful to Alain, Locardia and Sofche for their comments on some sections of this thesis, and Katrin and Stefen for their help with translating the summary into German. Thanks also to Tiemo Kahl, Alvaro Soares, David Forrester, Clara Arranz, Cristabel Duran Rangel, Masha van der Sande, Simone Ciuti, Somidh Saha, Rüdiger Unseld and Adam Benneter for helping me to clarify and find answers in times of uncertainty about statistical analysis. I would like to extend my thanks to Prof. Dr. Marielos Peña-Claros, Prof. Dr. Francis Putz and Prof. Dr. Helge Bruelheide for significant and important discussions that contributed to the improvement of some sections and to additional motivation for the development of this doctoral research. I also thank the Forest Ecology and Management group at Wageningen University for warmly receiving me for a short-term research mission, especially to Marielos, Masha and Lourens for discussing and sharing ideas and data on functional traits for the development of the third chapter of this thesis. In Freiburg, I felt very welcomed at my work group since the beginning to the very last moment. Thank you to all members of the “Waldbau-family” for that! Your friendly, respectful and lively atmosphere helped me feel at home and encouraged me to challenge myself, from exercising my German skills to confronting some beliefs. I thank Ursula Eggert, Bernardette Trautwein, Germar Csapek, Martin Kohler and Mathias Frowein for their prompt assistance throughout these years, and Prof. Dr. Albert Reif for always making the best for the group. Thanks for sharing good moments to Sofche, Juliane, Locardia, Cristabel, Peggy, Mario, Alvaro, Anderson, Fabiana, Adriana, Juanita, Katja, Rodrigo and Jan. Special thanks to my roommates Steffan Schneider and Fillipo Del Gatto for making the time at the office enjoyable and motivating, as well as to your wives and families for nice moments spent together.