Functional Diversity in the Hyper-Diverse Mangrove Communities in Papua New Guinea Lawong Balun [email protected]
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University of Tennessee, Knoxville Trace: Tennessee Research and Creative Exchange Doctoral Dissertations Graduate School 12-2011 Functional Diversity in the Hyper-diverse Mangrove Communities in Papua New Guinea Lawong Balun [email protected] Recommended Citation Balun, Lawong, "Functional Diversity in the Hyper-diverse Mangrove Communities in Papua New Guinea. " PhD diss., University of Tennessee, 2011. https://trace.tennessee.edu/utk_graddiss/1166 This Dissertation is brought to you for free and open access by the Graduate School at Trace: Tennessee Research and Creative Exchange. It has been accepted for inclusion in Doctoral Dissertations by an authorized administrator of Trace: Tennessee Research and Creative Exchange. For more information, please contact [email protected]. To the Graduate Council: I am submitting herewith a dissertation written by Lawong Balun entitled "Functional Diversity in the Hyper-diverse Mangrove Communities in Papua New Guinea." I have examined the final electronic copy of this dissertation for form and content and recommend that it be accepted in partial fulfillment of the requirements for the degree of Doctor of Philosophy, with a major in Ecology and Evolutionary Biology. Taylor Feild, Major Professor We have read this dissertation and recommend its acceptance: Edward Shilling, Joe Williams, Stan Wulschleger Accepted for the Council: Carolyn R. Hodges Vice Provost and Dean of the Graduate School (Original signatures are on file with official student records.) Functional Diversity in Hyper-diverse Mangrove Communities in Papua New Guinea A Dissertation Presented For the Doctor of Philosophy Degree The University of Tennessee, Knoxville Lawong Balun December 2011 Copyright © 2011 by Lawong Balun All rights reserved ii DEDICATION To my father, mother, father in law, mother in law Balun Garap, Marawe Garap, Lalai Kulang Yam, and Nang Gamun Yam and my wife Laden Kulang Balun iii ACKNOWLEDGEMENTS I thank Professor Taylor Feild, my advisor, for accepting me as his student and integrating my research into his NSF grant and for providing scholarly advice and critically reading the chapters of my dissertation. I also wish to express my gratitude to the members of my committee, Ed Schilling, Joe Williams, and Stan Wullschleger for providing me academic guidance and positive feedback. I thank Professor Gary McCracken (Head of the Department of Ecology and Evolutionary Biology, EEB) for the EEB Department financial support during my PhD tenure. I thank the course instructors of the EEB Core course: Professors Nathan Sanders (Coordinator), Daniel Simberloff, Jame Fordyce, Susan Riechert, Randall Small, Daren Husley, Benjamin Fitzpatrick, and Michael Gilchrist for providing in-depth insights regarding the core science of ecology and evolutionary biology. I thank several of my colleagues for their friendship, encouragement, and intellectual stimulation: Dr. MacKenzie Taylor, Angela Patino, Steve Furches, Annie Becker, Matt Valente, Michelle Smith, and Lisa Cantwell. I especially thank the past members of the Feild Lab, Dr. David Chatelet, Patrick Hudson, and Angela Patino for their lending hands in the anatomical laboratory setups. Field experiments and in situ data collection in the difficult working environment of the mangrove forest would not have been possible without the assistance of the following individuals and organizations in Papua New Guinea: University of Papua New Guinea iv Biology Department, particularly Dr. Augustine Mungkaje (Director, Motupore Island Research Center); Roga Gabiobu, and the security personals of the Motupore Research Center; Papua New Guinea University of Technology Forestry Department, particularly, Eko Maiguo (Acting Head), Trevor Galgal, and Jimmy Nona; New Guinea Binatang Research Center, particulary, Dr. Vojtech Novitney and Sam Legi; PNG Forest Research Institute, particularly Robert Kiapranis (Biology program leader), Billy Bau, Kipro Damas; and members of the Kulang family (Kenneth Kulang, Jeremiah Kulang, and Dorothy Kulang), staff of WWF-PNG office (Olo Gebia, Saina Jeffrey, and Roy Banka). I would like to express my deep gratitude to the Bosimbi family, Bidang family, Dorothy Kulang, and Kiapranis family in Papua New Guinea whose logistical support eased my field work. I especially thank Netesha Kulang Bidang for preparing nourishing meals for my team in the field. The following sources provided funding for my dissertation work: The National Science Foundation; PNG University of Technology; the University of Tennessee (Department of Ecology and Evolutionary Biology Summer Research Award [2009, 2010]). Finally, I would like to thank my wife, my children, grandson, in-laws, sisters, brothers, nieces and nephews, for their love and encouragement, which pushed me through this long journey. v ABSTRACT Variation in plant functional traits reflect the differences in an environment a species occupies and the variation in the functional traits across an environmental gradient and growth form reflects the functional performance of the coexisting species. Therefore, detecting the differences in functional traits of species is important to our understanding of the performance of species. The leaf functional traits; Leaf mass per area (LMA) and vein density (DV) are key traits in the resource use strategies and photosynthetic gas exchange capacity (Amax, gsmax) of all plants. Mangroves occupy a hypersaline narrow ecological range and therefore are thought to have uniform functional performance. This view makes sense for low diversity mangrove communities, but what about the hyperdiversity communities such as those in New Guinea? A comprehensive review of the present understanding on the ecology and socio-ecological values of mangrove was undertaken to establish a good foundational understanding of my study on New Guinea mangroves (Chapter 1). It is widely held that mangroves thriving in a hypersaline environment experience high water constraints and as a result have high water use efficiency, therefore have narrow functional performance. I investigated the leaf and photosynthetic functional traits of 31 mangrove species among different zonation bands and growth from six mangrove communities in New Guinea to test the view that mangroves have a narrow functional performance (Chapter 2). I then investigated the differences in wood and stem hydraulic traits among different zonation bands and growth forms and the relationship between wood hydraulics and leaf photosynthetic gas assimilation functional performance to further test the long standing view that mangroves have a narrow functional performance (Chapter 3). I then summarized the major findings of my two studies and their implications vi on mangrove restoration and rehabilitation with particular reference to the recent mangrove rehabilitation initiatives in Papua New Guinea (Chapter 4). My studies on leaf and wood functional traits across different zonation bands, growth forms and root system types consistently revealed that mangroves have a wide functional performance, and different species exhibited different resource use strategies. vii TABLE OF CONTENTS Chapter Page CHAPTER I: MANGROVE AND MANGROVE FOREST ..................................................... 1 INTRODUCTION ...................................................................................................................... 1 Mangrove global distribution and diversity ............................................................................ 6 The global significance of mangrove forests ...................................................................... 8 New Guinea mangroves ........................................................................................................ 18 Ecology of mangroves .......................................................................................................... 24 Mangrove zonation ........................................................................................................... 24 Forest structure.................................................................................................................. 29 Mangrove succession ........................................................................................................ 30 Salt Excretion .................................................................................................................... 33 Dissertation Focus: Understanding ecophysiological diversity in hyperdiverse mangrove communities in PNG ............................................................................................................. 42 LITERATURE CITED ............................................................................................................. 47 CHAPTER II: DIVERSITY OF LEAF PRODUCTIVITY FUNCTIONA FROM HYPERDIVERSE NEW GUINEA MANGROVE ECOSYSTEM ......................................... 75 Abstract ..................................................................................................................................... 76 INTRODUCTION .................................................................................................................... 78 LITERATURE CITED ........................................................................................................... 156 APPENDICES ........................................................................................................................ 172 CHAPTER III: WOOD MECHANICAL-HYDRAULIC FUNCTION ACROSS 31 SPECIES OF NEW GUINEA MANGROVES ......................................................................................