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Seed Rain and Seed Predation in a Hawaiian Dry Forest Community

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Seed Rain and Seed Predation in a Hawaiian Dry Forest Community INVESTIGATING SEED DISPERSAL AND SEED PREDATION IN A HAWAIIAN DRY FOREST COMMUNITY IMPLICATIONS FOR CONSERVATION AND MANAGEMENT A THESIS SUBMITTED TO THE GRADUATE DIVISION OF THE UNIVERSITY OF HAWAI‘I IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE IN BOTANICAL SCIENCES (BOTANY – ECOLOGY, EVOLUTION, AND CONSERVATION BIOLOGY) DECEMBER 2004 By Charles G. Chimera Thesis Committee: Donald Drake, Chairperson Gerald Carr David Duffy Lloyd L. Loope We certify that we have read this thesis and that, in our opinion, it is satisfactory in scope and quality as a thesis for the degree of Master of Science in Botanical Sciences (Botany – Ecology, Evolution and Conservation Biology). THESIS COMMITTEE Chairperson ii ACKNOWLEDGEMENTS I would like to thank the EECB Program, the Hawai‘i Audubon Society, the Dai Ho Chun Fellowship, and the Charles H. Lamoureux Fellowship in Plant Conservation for providing funding for my research. I would like to thank Sumner Erdman and Tony Durso of Ulupalakua Ranch for facilitating access to my study site through ranch lands. I would like to thank the staff of the Natural Area Reserve System, particularly Betsy Gagne, Bill Evanson and Bryon Stevens, for providing access to the Kanaio Natural Area Reserve as well as for logistical support. Lloyd Loope and Art Medeiros were extremely generous with their knowledge and guidance, factors that greatly influenced my decision to conduct research in Hawaiian dry forests. Lloyd Loope also provided vehicle support during the initial stages of my research. I thank my thesis committee chair, Don Drake, and my committee members, Gerry Carr, Dave Duffy and Lloyd Loope for providing me with guidance and assistance during all stages of my thesis research. Finally, I would especially like to thank my wife, Melissa, for providing me with unlimited patience and support during my entire graduate experience. iii TABLE OF CONTENTS ACKNOWLEDGEMENTS.............................................................................................. III LIST OF TABLES............................................................................................................ VI LIST OF FIGURES ....................................................................................................... VIII CHAPTER 1. INTRODUCTION ....................................................................................... 1 Loss of Native Seed Dispersers .................................................................................. 4 Rodents and Seed Predation........................................................................................ 5 Influence of Vegetation on Seed Dispersal and Predation.......................................... 6 Hypotheses.................................................................................................................. 8 CHAPTER 2. PATTERNS OF SEED DISPERSAL: VEGETATION STRUCTURE, FRUGIVORY AND SIZE EFFECTS .............................................................................. 10 Introduction................................................................................................................... 10 Loss of Native Seed Dispersers ................................................................................ 12 Influence of Vegetation on Seed Dispersal............................................................... 13 Methods......................................................................................................................... 15 Study Site.................................................................................................................. 15 Measurement of Seed Rain Under Trees Versus in Exposed Areas......................... 16 Seed Dispersal Analysis............................................................................................ 19 Bird Observations ..................................................................................................... 20 Seed Size Measurements........................................................................................... 21 Characterization of the vegetation ............................................................................ 22 Results........................................................................................................................... 22 Seed Rain .................................................................................................................. 22 Bird Observations ..................................................................................................... 33 Seed Size and Bird Dispersal.................................................................................... 35 Discussion..................................................................................................................... 36 CHAPTER 3: POST-DISPERSAL SEED PREDATION: LOCATION AND EFFECT ON SPECIES .................................................................................................................... 46 Introduction................................................................................................................... 46 Methods......................................................................................................................... 49 Study Site.................................................................................................................. 49 Measuring Seed Predation Levels............................................................................. 50 Treatments................................................................................................................. 51 Analysis..................................................................................................................... 53 Results........................................................................................................................... 54 Discussion..................................................................................................................... 62 iv CHAPTER 4: CONCLUSION ......................................................................................... 68 APPENDIX A. TREE DIMENSIONS (BASAL DIAMETER, HEIGHT, CROWN DIAMETER) OF STUDY SPECIES AND DEAD TREES............................................. 73 APPENDIX B. PHENOLOGY OF STUDY TREES AND COMMON FLESHY- FRUITED SHRUBS OF THE KANAIO NATURAL AREA RESERVE....................... 75 Methods......................................................................................................................... 75 APPENDIX C. RELATIVE FREQUENCY AND ABUNDANCE OF NON-NATIVE BIRDS IN KNAR STUDY SITE ..................................................................................... 82 Methods......................................................................................................................... 82 APPENDIX D. CHARACTERIZATION OF THE VEGETATION OF KNAR STUDY SITE .................................................................................................................................. 87 Methods......................................................................................................................... 87 LITERATURE CITED ..................................................................................................... 91 v LIST OF TABLES Table 2.1. Mean seed density by dispersal vector and location at KNAR from April 2003 to March 2004........................................................................................................... 23 Table 2.2. Mean density under trees and in exposed sites, and percentage of total seed rain for all species collected in 180 seed traps in KNAR from April 2003 to March 2004........................................................................................................................... 24 Table 2.3. Mean species richness per trap of fleshy-fruited seed rain under trees and in exposed sites for six dry forest trees......................................................................... 25 Table 2.4. Mean density of seeds dispersed by birds and fleshy-fruited seed rain under trees and in associated exposed sites for six dry forest trees in KNAR.................... 27 Table 2.5. Mean density of seeds dispersed by birds for 11 species under six dry forest trees in KNAR........................................................................................................... 28 Table 2.6. Relative frequency of fleshy-fruited species in the seed rain under six dry forest trees and all exposed seed traps at KNAR...................................................... 29 Table 2.7. Frequency of seedlings of fleshy-fruited species under six dry forest trees at KNAR. ...................................................................................................................... 33 Table 2.8. Mean visits per hour and mean fruits consumed per foraging visit by seven non-native bird species to five dry forest trees at KNAR......................................... 35 Table 2.9. Mean seed length and width of 14 seeds collected in the seed rain at KNAR.. ................................................................................................................................... 36 Table 3.1. Means of seed length and width and mean removal rate under trees and in exposed sites for species used in the seed predation trials........................................ 55 Table 3.2. General linear models for number of seeds and/or fruits remaining versus block, treatment, location,
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