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Phd Thesis Gerhard Overbeck

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Lehrstuhl für Vegetationsökologie Department für Ökologie Technische Universität München Effect of fire on vegetation dynamics and plant types in subtropical grassland in southern Brazil Gerhard Overbeck Vollständiger Abdruck der von der Fakultät Wissenschaftszentrum Weihenstephan der Technischen Universität München zur Erlangung des akademischen Grades eines Doktors der Naturwissenschaften (Dr.rer.nat.) genehmigten Dissertation. Vorsitzender: Univ.-Prof. Dr.rer.nat. Ludwig Trepl Prüfer der Dissertation: 1. Univ.-Prof. Dr.rer.nat. Jörg Pfadenhauer 2. Univ.-Prof. Dr.sc.techn. (ETH Zürich) Johannes Schnyder 3. Prof. Valério De Patta Pillar, Ph.D. (UWO/Canada), Universidade Federal do Rio Grande do Sul, Porto Alegre, Brasilien Die Dissertation wurde am 19.05.2005 bei der Technischen Universität München eingereicht und durch die Fakultät Wissenschaftszentrum Weihenstephan am 23.06.2005 angenommen. Contents Acknowledgements ................................................................................................................................. 2 Preface..................................................................................................................................................... 4 INTRODUCTION: Grasslands in southern Brazil – the setting............................................................. 5 CHAPTER 1: Floristic composition, environmental variation and species distribution patterns in burned grassland in southern Brazil with Sandra Cristina Müller, Valério DePatta Pillar & Jörg Pfadenhauer......................................... 17 CHAPTER 2: Small-scale dynamics after fire in South Brazilian humid subtropical grassland with Sandra Cristina Müller, Valério DePatta Pillar & Jörg Pfadenhauer......................................... 39 CHAPTER 3: Adaptive strategies to fire in subtropical grassland in southern Brazil with Jörg Pfadenhauer.......................................................................................................................... 62 CHAPTER 4: Ecosystem effects and responses: trait-based analysis of plant functional types in burned grassland with Sandra Cristina Müller, Valério DePatta Pillar & Jörg Pfadenhauer......................................... 91 CHAPTER 5: No stimulation of germination by heat found in subtropical grassland species with Sandra Cristina Müller, Valério DePatta Pillar & Jörg Pfadenhauer....................................... 110 CHAPTER 6 (Conclusion): Fire in grassland – disturbance or management tool?............................ 121 SUMMARY ........................................................................................................................................ 131 ZUSAMMENFASSUNG.................................................................................................................... 133 RESUMO ............................................................................................................................................ 135 1 Acknowledgements Principally, I would like to thank Prof. Dr. Jörg Pfadenhauer for the possibility to conduct this work, for the always constructive and motivating discussions and for freely sharing all his experience with me. Likewise, I would like to express my thanks to Prof. Dr. Valério DePatta Pillar, who „adopted“ me in his lab as one of his students, supported my work in numerous ways and always found time for discussions. My great and special thanks goes to my colleague Sandra Cristina Müller, who not only introduced me to the vegetation of southern Brazil, helped organizing my work and stay in Brazil in the beginning, but principally turned to be a great friend throughout and beyond this work – muitissimo obrigado mesmo! I would like to thank my colleagues in the Probral project, Andrea Zellhuber, Eduardo Días Forneck and Wolfram Adelmann, for all help, discussions and good times we had in Brazil or Germany. A great thanks also goes to Profa. Dra. Maria Luiza Porto, for organization of the Probral project, support of and advice on my work, and the possibilities to get a good overview of vegetation and landscapes of southern Brazil in an impressive excursion. Further thanks go to Profa. Dra. Sandra Hartz, coordinator of the PPG Ecologia of the UFRGS, for all support of my project and all administrative help during my stays in Brazil. Dankeschön also to Frau Tork, who managed to resolve many organisatorial matters in a uncomplicated way, despite the distance between Freising and Porto Alegre. Obrigado also to all the friends and colleagues at “Ecoqua” and the UFRGS’ Department of Ecology, especially Carolina Blanco da Rosa, Claudia Beltrame Porto, Marcella Pinillos, Melina Grosellini dos Santos, Silene de Carvalho, Vanessa Guazulha Porto, Ênio Egon Sosinski Jr., Glauco Schüssler, José Pedro Pereira Trindade, Julia-Maria Hermann (another “invader” like me), Juliano Morales de Oliveira and Rafael Engelman Machado, with all of whom I had many good discussions on work of my thesis or related research, and, even more important, who introduced me to many great things, from how to prepare a chimarrão and how to do the rala-bucho to how to conduct a bootstrap resample. It was great to share these past three years with you all, e não vou esquecer o meu lado gaúcho! In different parts of my work, a number of person were of invaluable help. Principally, I would like to thank Alessandra Fidelis, Tina Schlossorsch, Sebastian Klebe and Pascual García Martinez for their participation in the overall project and their respective Master or Diploma thesis work, each of great value for my own thesis. Alessandra Fidelis deserves a special thanks for help in some of the experiments and for continuous discussion, in Brazil or in Germany, which I hope will 2 continue in future projects on grasslands in southern Brazil. Further, I would like to thank Emiliano Santarosa, Marcos Carpes Westfalen, Ana Luiza Leichter Matte and Márcia Ruff da Silva for assistance in some of the field work and experiments. I owe my thanks to staff at the UFRGS Department of Botany for help with identification of collected plant material, principally to Profa. Dra. Ilsi Iob Boldrini, Profa. Dr. Lillian Eggers, Profa. Dra. Hilda Maria Longhi-Wagner and Prof. Paulo Brack. Prof. Dr. Renato Borges de Medeiros gave valuable advice concerning seed bank and germination experiments, and Profa. Dra. Lucia Brandão Franke made it possible to work in the UFRGS Department of Agronomy’s seed lab. Additionally, I would like to thank staff at the Instituto do Meio Ambiente of the PUCRS, Porto Alegre, especially Prof. Ricardo Mello, for the possibility to work on Pró-Mata, and staff there for all the support. My thanks also go to UFRGS security staff and personnel from the Batalhão Ambiental of the Brigada Militar, for always guaranteeing our safety in the field. Adroaldo deserves a special Obrigado for sharing with us the secrets of the true cozinha campeira. I further would like to thank the German National Academic Foundation (Studienstiftung des Deutschen Volkes) for the PhD scholarship, DAAD and CAPES, cooperating under the ProBral program, for covering of travel expenses, and the DFG (German Research Foundation) for additional support of the project. Finally, I would like to thank Fabíola for her patience in times I was thinking too much of my work, for passing close to me even the times I was long away and for all support when I needed it most – obrigado! 3 Preface The presented thesis is written as a cumulative PhD thesis. Chapters 1 to 5 each are independent papers that have been submitted for publication in different scientific journals. A general introduction and a concluding chapter with an integrating discussion (Chapter 6) create the necessary frame for joint consideration of the different chapters. Each of these chapters (papers) originally had been formatted according to the instructions given by the respective journal. To facilitate reading of the thesis, tables and figures were always inserted into the text and formatted in the same way, and font size, line spacing and captions were adjusted to one format. Other formatted objects, e.g. of references in the text and in the bibliography were left unchanged, i.e. may differ slightly in style between the different chapters, depending on the rules of the journal where the paper was submitted. Enumeration of tables and figures was conducted separately for each chapter. In addition to the synthesizing summaries in English and German, the thesis at its end contains an expanded summary in Portuguese, briefly presenting the contents of each of the different chapters. 4 INTRODUCTION Grasslands in southern Brazil – the setting 5 Introduction: Grasslands in southern Brazil Overview of present vegetation and vegetation history of southern Brazil Due to its geographic position around the 30° parallel of latitude, a limit for tropical vegetation types (Cabrera & Willink 1973), and at the eastern side of South America, southern Brazil is characterized by a transitional situation between tropical and temperate climate, with hot summers and cool winters and no dry season. Differences in geological substrate and altitudinal variation further contribute to diversity of vegetation types in the region (Waechter 2002). The natural vegetation in Brazil’s southernmost state, Rio Grande do Sul (RS), presents itself as mosaics of grassland, shrubland and different forest types (e.g. IBGE 1986, Leite & Klein 1990, Pillar & Quadros 1997; see Fig. 1). B R A Z I L Atlantic forest Location of the
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    BALDUINIA. n. 22, p. 15-22, 15-V-201O ANATOMIA DAS MADEIRAS DE EUGENIA BURKARTIANA (D. LEGRAND) D. LEGRAND E MYRCIARIA CUSPIDATA O. BERG, DUAS MIRTOÍDEAS NATIVAS NO RIO GRANDE DO SUV SIDINEI RODRIGUES DOS SANTOS2 JOSÉ NEWTON CARDOSO MARCHIORI3 RESUMO No presente estudo são descritos e ilustrados os caracteres anatômicos do lenho de Eugenia burkartiana D. Legrand (D. Legrand) e Myrciaria cuspidata O. Berg, a partir de material coletado no Rio Grande do Sul. As características anatômicas em comum corroboram a conhecida homogeneidade estrutural das Myrtaceae. Entre os caracteres anatômicos diferenciais, salientam-se o arranjo do parênquima axial, a presença e/ou abundância de conteúdos nos raios e de séries cristalíferas no parênquima axial, bem como a composição celular nas margens de raios e a freqüência de poros. Palavras-chave: Eugenia burkartiana, Myrciaria cuspidata, anatomia da madeira, Myrtaceae. ABSTRACT [Wood anatomy of Eugenia burkartiana (D. Legrand) D. Legrand and Myrciaria cuspidata O. Berg, two native Myrtoideae from Rio Grande do Sul state, Brazil]. The wood anatomy of Eugenia burkartiana (D. Legrand) D. Legrand and Myrciaria cuspidata O. Berg are described and illustrated, based on samples from Rio Grande do Sul state, Brazil. The anatomical features shared by the two species reinforce the well known structural homogeneity ofMyrtaceae farnily. To segregate both species are specially important: the arrangement ofaxial parenchyma; the presence and/or abundance of organic inclusions in rays; the presence of crystalliferous strands on axial parenchyma; the cellular composition of ray margins (uniseriate ends); and the frequency of pores. Key words: Eugenia burkartiana, Myrciaria cuspidata, wood anatomy, Myrtaceae.