ANT COMMUNITIES OF FLORIDA’S UPLAND ECOSYSTEMS: ECOLOGY AND SAMPLING By JOSHUA R. KING A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY UNIVERSITY OF FLORIDA 2004 Copyright 2004 by Joshua R. King To my wife. Thank you for teaching me what is truly important in life. ACKNOWLEDGMENTS I would like to thank my committee members John Capinera, Mark Deyrup, Robert McSorley, Sanford Porter, and Kenneth Portier for reading the dissertation and providing sound advice on the politics of academia, statistics, publishing, teaching, and the pursuit of biological knowledge. Their contributions have all aided in my development as a scientist, collaborator, and colleague; for that I am grateful. In particular I would like to thank my advisor, Sanford Porter. None of this work could have been accomplished without the support, laboratory space, equipment, and encouragement he provided. I am grateful to Lloyd Davis for teaching me to be a better collector and observer of the natural world. A man who has forgotten more entomology than I will ever know, he has impressed on me that a broad entomological knowledge is the best context within which to build an understanding of ants. I am indebted to Mark Deyrup for showing me that it is possible to know how to identify everything, and that I must not forget that sampling and theory can never replace collecting and natural history. Thanks go to Lloyd Morrison for sharing ideas, insight on being a better scientist, surfing, and Frisbee. I thank Sanford Porter and Walter Tschinkel for sharing ideas and showing me the importance of a mechanistic, experimental approach to studying ants. I also thank Walter Tschinkel and his lab group for sharing ideas and being patient while I finished. I thank Lloyd Davis and Mark Deyrup for assisting with and verifying species identifications. I am also grateful for myrmecological advice from Stefan Cover, without iv which the project would not have been as successful. I thank the Archbold Biological Station and Mark Deyrup for laboratory space and accommodation during part of this work. I thank the University of Florida, the Florida Department of Environmental Protection’s State Parks Division, and the U.S. National Forest Service for permission to perform sampling in the Katherine Ordway Biological Preserve, San Felasco Hammock State Park, and Osceola National Forest, respectively. Voucher specimens from this project have been donated to Harvard’s Museum of Comparative Zoology and the Archbold Biological Station. I thank the University of Florida for financial support during part of my graduate studies in the form of a University of Florida Alumni Fellowship. I also thank Walter Tschinkel for financial support during the completion of the dissertation. I sincerely thank my mother, Pamela Snow, for instilling in me a love of scholarship, the natural world, and writing. I am also indebted to her for teaching me patience and the desire to always be optimistic and forward-moving, no matter how rough the going gets or how daunting the task. I am indebted to my wife, Kari, for teaching me to be more disciplined. I am also grateful to my wife for emotional support and for sharing her life with me. Without these things I would not have been able to complete the dissertation. Finally, I thank my daughter Maizie for showing me what a joy life can be. v TABLE OF CONTENTS page ACKNOWLEDGMENTS ................................................................................................. iv LIST OF TABLES........................................................................................................... viii LIST OF FIGURES ........................................................................................................... ix ABSTRACT.........................................................................................................................x CHAPTER 1 INTRODUCTION ........................................................................................................1 2 ASSEMBLY RULES FOR INSECTS AT LOCAL AND REGIONAL SCALES: ABUNDANCE, DIVERSITY, AND BIOMASS OF ANTS IN FLORIDA’S UPLAND ECOSYTEMS .............................................................................................4 Introduction...................................................................................................................4 Study Area and Methods ..............................................................................................8 Upland Ecosystems ...............................................................................................8 Inventory Design .................................................................................................12 Analysis ...............................................................................................................14 Results.........................................................................................................................22 Species Richness .................................................................................................22 Abundance and Biomass .....................................................................................22 Behavioral Dominance ........................................................................................27 Species Co-occurrence ........................................................................................28 Introduced Species...............................................................................................29 Discussion...................................................................................................................30 Taxocene Attributes.............................................................................................30 Biogeography, Synthesis, and Applications........................................................50 3 EVALUATION OF SAMPLING METHODS AND SPECIES RICHNESS ESTIMATORS FOR ANTS IN UPLAND ECOSYSTEMS IN FLORIDA..............78 Introduction.................................................................................................................78 Methods ......................................................................................................................83 Study Area...........................................................................................................83 Sampling..............................................................................................................83 vi Analysis ...............................................................................................................85 Results.........................................................................................................................90 Observed and Estimated Species Richness .........................................................90 Rarity ...................................................................................................................92 Complementarity of Ecosystems.........................................................................92 Effectiveness of Sampling Methods....................................................................93 Discussion...................................................................................................................95 Inventory Completeness ......................................................................................95 Performance of Species Richness Estimators......................................................97 Rarity ...................................................................................................................98 Efficiency of Sampling Methods.......................................................................100 4 CONCLUSION.........................................................................................................113 LIST OF REFERENCES.................................................................................................116 BIOGRAPHICAL SKETCH ...........................................................................................132 vii LIST OF TABLES Table page 2-1 Species list ................................................................................................................59 2-2 Species richness, slope (m), and R2 values for fitted lines .......................................63 2-3 Five most abundant species ......................................................................................64 2-4 Co-occurrence patterns of ants .................................................................................65 2-5 Body size overlap patterns of ants............................................................................66 2-6 Ant species turnover .................................................................................................67 3-1 Species richness estimates and measures of inventory completeness ....................105 3-2 Mean percent faunal complementarity among ecosystems ....................................106 3-3 Percent faunal complementarity of sample methods..............................................107 viii LIST OF FIGURES Figure page 2-1 Map of Florida..........................................................................................................68 2-2 Rarefaction curves for ecosystems ...........................................................................69 2-3 Relationship between species occurrences and body