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Forest Gaps Structure and Tree Regeneration in a Montane Guayanan Forest, Sierra De Lema, Venezuela

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Forest Gaps Structure and Tree Regeneration in a Montane Guayanan Forest, Sierra De Lema, Venezuela Forest gaps structure and tree regeneration in a montane Guayanan forest, Sierra de Lema, Venezuela Thesis submitted in partial fulfilment of the requirements of the degree of Dr. rer. nat of the Faculty of Forest and Environmental Sciences Albert-Ludwigs Universität, Freiburg im Breisgau, Germany by Cristabel Durán Rangel Freiburg im Breisgau, July 2011 Dean: Prof. Dr. Juergen Bauhus First advisor: Prof. Dr. Dr. h.c. Albert Reif Second reviewer: Prof. Dr. Siegfried Fink Disputation term: 29 November 2011 for Aida, Haiddye and Moritz for the majestics Tepuys and breathtaking Gran Sabana Acknowledgments ªcaminante no hay camino, se hace camino al andarº ªwanderer, there is no path, the path is made by walkingº Antonio Machado (1875 ± 1983), Poet. This is the longest hike I ever wandered to reach the summit. The person who accompanied me from the planning of the journey to the end of this hike, is the person I most have to thank: mein lieber Doktorvater the Prof. Albert Reif. Thank you very much for your enormous support in all senses! I have been very lucky to meet you and share with you these years! My second adviser the Prof. Lionel Hernández introduce me in the scientific world and in the tropical ecology, I thank all your advice and understanding. I thank the Prof. Siegfried Fink for be my second reviewer. Several institutions made this study possible. I thank to Prof. Juergen Bauhus, as the representer of the Silviculture Institute, for permit me to work here for a long time. The Centro de Investigaciones Ecológicas de Guayana, from the Universidad Nacional Experimental de Guayana provided me the institutional and logistic support in Venezuela. I also thank the Prof. Pelz and the Institute of Biometry for be working in their offices in the time of renovations at our institute. The Institute for National Parks in Venezuela, Petróleos de Venezuela and Parupa Research Station in Gran Sabana provided necessary logistical support for this study. This study was partially financed by several institutions: the State Electricity Enterprise EDELCA C.A., in Venezuela and the International Promotions Program (IPP) of the Faculty of Forestry and Environmental Sciences at the University of Freiburg. The IPP paid the language correction of the thesis and scientific articles. The Wissenschaftliche Gesellschaft of the University of Freiburg supported this work funding my assistance to congresses in Venezuela. I received financial support through grants by the Verband der Freunde der Universitaet Freiburg, the Dr. Leo-Ricker-Stiftung in Freiburg im Breisgau and the Landesgraduiertenfoerderung, Baden-Wuerttemberg Ministry of Science, Research and Art, Germany. A very specially thank goes for Stefanie Gaertner. You give me so much scientific input and understanding. We are happy to have you again between us! I thank the Prof. Miguel Acevedo for the time I spend improving my English and analyzing hemispherical pictures at the University of North Texas. You introduced me in the powerful and exiting world of R! The botanical identification was possible with the collaboration of Elio Sanoja, Roland Keller and Bruce Holst. Several person give advices during the realization of this study: Florian Straub, Peter van der Meer, David Butler Manning. Katja Seis translated the summary of this work in German. Nestor Gutiérrez checked the summary in Spanish and improved the format of the final document. A special thanks goes for the Dr. Sibel Vurgun for her effectively and valorously advising related to time management, thesis structure and self-confidence during the last phase of my work! i I also thank to Markus Heck and the members of the GAUS-Group at the Fraunhofer-ISE for their understanding and flexibility in the time before the submission of my dissertation. One of the biggest profits I had in this time was the learning of the Open Source Operation Systems and Softwares, specially Linux and R. I appreciate the encourage and discussions with the R-Linux Waldbau-Community, namely: Dimitris and Katja! I felt very comfortable at the Institute of Silviculture, this is because all the Institute members are very nice and easy-going. I got help and exiting moments in field from colleagues and friends: Haiddye Durán, Rosita Pabón, Moritz Moessner, Leandro Salazar, Jesús Salazar, Marc Urlich, Seth Kaupinnen, Philipp Loeper, Jackeline Ortíz, Anselmo, Patricia Rodriguez, Mayerslin Echagaray, José Ayala, Elio Sanoja, Roland Keller, Hernán Castellanos and Luz Delgado. Last by not the least my dear friends and families! All you make life more beautiful and enjoyable! Thanks for all the nice moments with you: Osvaldo, Silvio, Marianna, Chunling, Rodrigo, Jan and Hendrik. A very special place in my heart is deserved for: Nestor, Maria Dolores, Dimitris, Juanita and Katja. You accompanied me in the good and difficult moments! During my time in Germany I won not one, but two loving families! In Freiburg: Nelida, Silvana, Johanna and Albert. In Winnenden-Heilbronn: Joerg, Barbara and Ingeborg. Thank you for be there when I needed. Without the support and sacrifices of my mother Aida, my most important achievements will be impossible. My sister Haiddye and Miguel Angel supported me during this time without critics or demands. (Sin el apoyo y los sacrificios de mi mamá Aida, mis más importantes logros habrian sido imposible. Mi hermana Haiddye y Miguel Angel me apoyaron durante éste tiempo sin críticas o exigencias. Los quiero muchísimo!). A very special place in this acknowledgements is deserved to my dear Moritz, my ªSeilschaftpartnerº: You made this large hike lighter, sunny, exiting and beautiful! MUCHAS GRACIAS A TODOS! Cristabel ii Table of Contents Acknowledgments...................................................................................................i 1 Introduction.........................................................................................................1 1.1 The role of gaps in tropical forests......................................................................1 1.2 Relevance of the research....................................................................................3 1.3 Objectives...........................................................................................................4 2 Study area............................................................................................................7 2.1 Location..............................................................................................................7 2.2 Physiography......................................................................................................7 2.3 Geology and geomorphology...............................................................................8 2.4 Soils....................................................................................................................9 2.5 Climate.............................................................................................................10 2.6 Vegetation.........................................................................................................10 3 Research undertaken.........................................................................................13 3.1 Understanding small-scale disturbances in Guayana©s montane forests: gap characterisation in the Sierra de Lema, Venezuela...........................................13 3.1.1 Objectives....................................................................................................13 3.1.2 Methods......................................................................................................14 3.1.3 Principal findings........................................................................................15 3.1.4 Discussion...................................................................................................16 3.1.5 Conclusions.................................................................................................17 3.1.6 Contribution of the authors ........................................................................17 3.1.7 Reference....................................................................................................17 3.2 Forest gaps without pioneers? Species composition and ecological groups within tree regeneration in a montane forest, Venezuelan Guayana...........................18 3.2.1 Objectives....................................................................................................19 3.2.2 Methods......................................................................................................19 3.2.3 Principal findings........................................................................................21 3.2.4 Discussion...................................................................................................24 3.2.5 Contribution of the authors.........................................................................26 3.2.6 Reference....................................................................................................26 3.3 Do microhabitats in forest gaps influence the tree regeneration? A study in the montane forests of the Venezuelan Guayana...................................................26 3.3.1 Objectives....................................................................................................27 iii 3.3.2 Methods......................................................................................................27 3.3.3 Principal findings........................................................................................29
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