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Natural and Experimental Tree Establishment in a Fragmented Forest, Ambohitantely Forest Reserve, Madagascar

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Natural and Experimental Tree Establishment in a Fragmented Forest, Ambohitantely Forest Reserve, Madagascar Natural and Experimental Tree Establishment in a Fragmented Forest, Ambohitantely Forest Reserve, Madagascar INGAR PARELIUSSEN DR. SCIENT THESIS Department of Biology Faculty of Natural Sciences and Technology Norwegian University of Science and Technology 2004 2004:14 ISSN 1503-8181 ISBN 82-471-6228-8 (Electronic) ISBN 82-471-6229-6 (Printed) Acknowledgments I would like to thank my supervisors, E. Gunilla Almered Olsson and W. Scott Armbruster for their continuing help through out the project. I would also like to thank Mary Edwards, Nancy Bazilchuk, Richard Strimbeck, Olga Hilmo, Terje Thun and Carolyn Baggerud for comments on the thesis. I would like to thank our collaborators in the Madagascar, the staff at ESSA-Forêt and ANGAP, especially Joel Ratsirarson, ESSA and Richard Ranarisoa ANGAP. A special thank goes to Jean Jacques Rasolofonirina, your knowledge of the study site and the plants in Ambohitantely were invaluable. I would also like to thank the Norwegian and Malagasy students that accompany me in field: Monica Lundberg, Sølvi Wehn, Susanne Hanssen and Sitraka Rakotoson. It has been a delight to work along some extraordinary helpful and inspiring Ph.D. students: Linda Dalen, Dagmar Hagen, Kristian Hassel, David Rae and Bård Solberg. I would also like to thank some of the faculty for answering questions and lending me books and ears: Marit By, Urban Gunnarsson, Håkan Hytteborn, Sigrid Lindmo, Johanne Maad, Bård Pedersen and Lars Söderström. When the writing block has been unsurmountable or the Ph.D. blues have festered, I have been blessed with many helpful and supportive friends which I give my deepest expression of gratitude to, you know who you are. My family have always been there for me and without their support I would not have written any thesis, ever. I would like to thank NFR for the research grant, and finally Magne Mæhre for the support he has given me all these years. III Papers 1. Ingar Pareliussen, E. Gunilla A. Olsson, Sølvi Wehn, Joel Ratsirarson. Natural establish- ment of trees in the grasslands between forest fragments in upland Madagascar. (Manuscript) 2. Ingar Pareliussen, E. Gunilla A. Olsson, W. Scott Armbruster. Factors limiting the survival of native tree seedlings used in conservation efforts at the edges of forest fragments in upland Madagascar. (Accepted for publication in Restoration Ecology) 3. Ingar Pareliussen, W. Scott Armbruster, E. Gunilla A. Olsson, Mary Edwards, Joel Ratsir- arson. Factors limiting the growth of native tree seedlings at the edges of forest fragments in upland Madagascar. (Manuscript) 4. Sølvi Wehn, E. Gunilla A. Olsson, and Ingar Pareliussen. Natural recovery of trees after fire in the edges of forest fragments in Ambohitantely forest, upland Madagascar. (Submitted) IV Contents Acknowledgments . III Papers . IV General introduction 9 Introduction . 9 Roads to deforestation and fragmentation . 9 Ecological effects of deforestation and fragmentation . 10 Challenges in reforestation of grasslands . 12 Aims . 16 Study area . 17 Madagascar . 17 Ambohitantely . 17 Selection of study sites . 20 Selection of species used in the experiments . 20 The sub-studies . 21 The grass sward as a barrier to tree establishment . 22 Micro-climate . 23 Soil factors . 24 Lack of tree seeds in the grassland . 25 Fire succession . 27 Is there continuing fragmentation of Ambohitantely forest? . 28 Forest fragmentation and its effect on tree reproduction . 28 Main findings . 30 Future management . 30 Future studies . 31 Literature cited . 41 Paper I 41 Natural establishment 43 Introduction . 43 Study area . 44 V Methods . 45 Selection of sites . 45 Species and dispersal modes . 45 Statistical considerations . 47 Results . 47 Species abundance and composition . 47 Comparing the grassland with the forest . 51 Turnover . 51 Growth . 51 Discussion . 53 Conclusions . 55 Acknowledgments . 55 Paper II 59 Factors limiting the survival of native tree seedlings 61 Introduction . 61 Methods . 62 Study area . 62 Selection of sites . 63 Selection of tree species . 63 Execution of experiments . 65 Statistical analysis . 66 Results . 66 The effect of site . 66 The effect of the experimental treatments . 67 The effect of the cover of the naturally establishing shrubs . 68 Discussion . 69 Distance from the forest . 69 Application of soil from the forest . 70 Fertilization . 71 Shading . 71 Soil moisture . 71 Importance of site . 71 Conclusion . 72 Acknowledgments . 72 VI Paper III 77 Factors limiting the growth of native tree seedlings 79 Introduction . 79 Methods . 80 Study area . 80 Site selection . 81 Tree species used in the experiment . 82 The experimental factors . 82 Execution of experiments . 83 Statistical analysis . 84 Results . 84 Sites . 84 Species . 84 Distance . 86 Forest soil . 86 Fertilizer . 88 Shading . 88 The naturally establishing shrub cover . 88 Discussion . 88 Main findings . 88 Comparison of growth and survival patterns . 90 The factors . 90 The effect of moisture . 91 Conclusion . 92 Acknowledgments . 92 Paper IV 97 Natural recovery of trees after fire 99 Introduction . 100 Fire . 100 Madagascar . ..
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