© 2008 James A. Macdonald ALL RIGHTS RESERVED
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© 2008 James A. MacDonald ALL RIGHTS RESERVED VARIATION AMONG MANGROVE FORESTS AS FISH HABITAT: THE ROLE OF PROP-ROOT EPIBIONTS, EDGE EFFECTS AND BEHAVIOR IN NEOTROPICAL MANGROVES by JAMES A. MACDONALD A Dissertation submitted to the Graduate School-New Brunswick Rutgers, The State University of New Jersey In partial fulfilment of the requirements for the degree of Doctor of Philosophy Graduate Program in Ecology and Evolution Written under the direction of Judith S. Weis, Ph.D. And approved by Judith S. Weis, Ph.D. Rebecca Jordan, Ph.D Peter Morin, Ph.D. Joseph Rachlin, Ph.D. New Brunswick, NJ May 2008 ABSTRACT OF THE DISSERTATION VARIATION AMONG MANGROVE FORESTS AS FISH HABITAT: THE ROLE OF PROP-ROOT EPIBIONTS, EDGE EFFECTS AND BEHAVIOR IN NEOTROPICAL MANGROVES by James A. MacDonald Dissertation Director: Judith S. Weis, Ph.D. Mangrove forests are an important nursery habitat for many species of reef fish, as well as a key component of the interlinked mangrove-seagrass-reef system. However, understanding how juvenile fish utilize the mangrove habitats is hampered by variation between mangrove habitats. This study sought to examine some reasons for variation between mangroves, focusing on physical characteristics, particularly the influence of sessile epibiont organisms. Using visual census, fish and sessile communities were compared in Rhizophora mangle roots in Bocas Del Toro, Panama, and Utila, Honduras. The results revealed significant positive correlation between depth, epibiont diversity, and density and fish species diversity and biomass. In order to determine a causal relationship between epibionts and fish community variation, two field experiments were established. In one, artificial mangrove roots (AMR) with different sets of artificial (AE) or real epibionts were established in five different locations. In the second experiment, fish were surveyed in 12 different mangrove transects, epibionts were reduced in half of those transects, and then surveyed again. In the artificial mangrove plots, treatments with the most heterogeneous structure had the greatest abundance and diversity of fish. When epibionts were reduced, fish ii abundance went up and biomass stayed level in controls, but abundance stayed flat and biomass decreased in treatment transects. The data indicate that epibionts can enhance fish abundance and diversity in mangroves, although the relationship may depend upon the specific epibionts. Separately, a series of prop-roots were surveyed and placed inside predator exclusion cages. After three months, the cages were removed. The results suggested that grazing mostly does not impact prop-root epibiont coverage. In a separate study, fish and epibiont communities were measured along a gradient within the mangroves away from boundaries of the mangrove forest. Both fish diversity and abundance showed a significant linear decrease away from the edge. Average size decreased as well, but epibionts showed no significant changes along the gradient. Last, juvenile reef fishes were captured by in mangroves and seagrass, tagged, and monitored for three months. All recaptures were within 10 meters of the original capture, but most occurred within a short time. iii ACKNOWLEDGEMENTS: An undertaking of this complexity could not be completed without the support and assistance of innumerable people along the way. The most obvious person to thank is of course my major professor, Judy Weis, who has read far too many drafts, seen too many powerpoints, and read far too many angry dispatches from various swamps for her own good and sanity. The fact that she still provided such valuable input despite such a ceaseless onslaught is a testament to her abilities as a scientist and a mentor, and I am eternally grateful for her assistance and support. I also want to thank my committee members, Rebecca Jordan, Peter Morin, and Joseph Raclin, whose comments, ideas, and sympathetic ears, despite recalcitrant mail servers were invaluable. Peter Morin’s forced proposal writing stands out particularly as an indispensable exercise. I also had invaluable support from colleagues in and around Rutgers, particularly from the faculty, staff, cast and crew of the Graduate Program in Ecology and evolution, particularly Marsha Morin, who has kept us all going at one time or another. Then there are friends, both in New Brunswick and Newark, particularly Jenn Adams-Krummins, Shannon Galbraith, Dominic D’Amore, Jane Zagajeski, Jessica Schnell, Cristy Cincotta, and Darshan Desai. I would like to especially thank Claus Holzapfel for putting up with my incessant statistics questions, many of them weird, and for generally being a great guy. In my own lab, I couldn’t have functioned without the advice of Terry Malcolm, Tish Robertson, Lauren Bergey, Allie Candelmo, and especially Jessica Reichmuth, my lab- mate and partner in crime since the beginning. I also want to single out two students, iv Ross Roudez and Suzan Shahrestani, who both helped me keep my projects going through thick and thin, especially Suzan, who took the brunt of the thick part, much of which is still clinging to her shoes. I also had help from a colorful cast of characters encountered throughout the American tropics. In Panama, I particularly want to thank Rachel Collin, Gabriel Jacome, Plinio Gondola, Steve Barnes, Patrick Irwin, Elise Granek, Scott Walls, Allie, Justin, Saray Dugan, Tito Lawrence (true, he was a thief, but he helped me a lot and I learned even more), and last but not least, the developer of Sunset Point, who is so awful that he inspired me anew to keep fighting, especially against him. In Honduras, a special thanks to the numerous people who sacrificed part of their Caribbean dive vacation to wade in dangerous swamps with me: Eleri Varney, Ben Mongin, Laura Astorquiza, Eric Topham, Nick Suarez, Steve Lawton, Linda Burt, Paddy Denning, James Saunders, Paula Doria, Ginny Brady, Ori Friedland (partner on the Utila death march), Andy Avery, Charli Todd, Isaac Lichtenstein, Dave the teacher, Callie, CJ, Ms. Tanya, goldtooth, and the snake. Last, I want to thank my family, my mother and father Judith and George (who would have approved, I’m sure) MacDonald and my sister Elizabeth MacDonald, for putting up with my interest in all things creepy, crawly, and far away since I was old enough to crawl, as well as my friends, Niall Dunne, Carlos “trans-Australian odyssey” Dedesma, Robert Schelly, and Ian Harrison, the latter two of which were happy to drop everything to dissect a fish in the kitchen. v Most importantly, I need to thank my wife, Sunny Hwang, who, among other things, put up with my long absences, kept me sane, and is generally far more understanding and wonderful than I ever deserved. It goes without saying that none of it would have been possible without generous support from funders: The Smithsonian Tropical Research Institute, The Academic Excellence Fund of the Graduate Program in Ecology and Evolution, Rutgers New Brunswick, Operation Wallacea, The PADI Foundation, The Rutgers Marine Field Station. vi TABLE OF CONTENTS ABSTRACT OF DISSERTATION ...........................................................................................ii ACKNOWLEDGEMENTS ....................................................................................................iv LIST OF TABLES .............................................................................................................xii LIST OF FIGURES ...........................................................................................................xiv CHAPTER I: INTRODUCTION ...........................................................................................1 I.1: HYPOTHESES ..................................................................................................9 CHAPTER II: REAL-WORLD CORRELATIONS OF EPIBIONTS WITH DIVERSITY, BIOMASS, AND COMMUNITY STRUCTURE OF FISHES ................................................................ 13 INTRODUCTION......................................................................................................... 13 MATERIALS AND METHODS ...................................................................................... 15 II.1: STUDY SITES ............................................................................................... 15 II.2: FISH SURVEYS............................................................................................. 19 II.3: EPIBIONT SURVEYS ..................................................................................... 20 II.4: ABIOTIC FACTORS ....................................................................................... 21 II.5: STATISTICAL ANALYSIS............................................................................... 21 RESULTS .................................................................................................................. 22 II.6: GENERAL RESULTS ..................................................................................... 22 II.7: DIVERSITY AND ABUNDANCE OF FISHES ...................................................... 23 II.8: AGE CLASSES.............................................................................................. 28 II.9: ABIOTIC FACTORS ....................................................................................... 29 II.10: COMMUNITY LEVEL EFFECTS ....................................................................30 vii DISCUSSION ............................................................................................................