METAPOPULATION DYNAMICS AND LANDSCAPE ECOLOGY OF THE FLORIDA SCRUB-JAY, APHELOCOMA COERULESCENS By BRADLEY M. STITH 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 1999 Dedicated to my wife and best friend, Ellen Mary Thorns ACKNOWLEDGMENTS I wish to acknowledge each of my committee members for their unique contributions to this dissertation. My chairman, Dr. Stephen R. Humphrey, stimulated my interests in issues relating to philosophy, policy, and management. His broad interests and ability to take on multiple careers were remarkable. In my times of need he provided unfailing support while allowing me great freedom to pursue my own course of study. As director of Archbold Biological Station, co-chairman Dr. John W. Fitzpatrick provided me with a wonderful opportunity to study the Florida Scrub-Jay at this world-class research facility. His ability to perform brilliantly as director, researcher, and conservationist is inspirational. I am grateful to him for sending a generous U.S.F.W.S. grant my way to support me towards the end of my program. I was also privileged to work under the tutelage of the legendary pioneer of Florida Scrub-Jay research, Dr. Glen E. Woolfenden, who showed me the world from a Florida Scrub-Jay's perspective. My collaborative research with Dr. Lyn Branch on the Vizcacha and the Florida scrub lizard, was a joy. She forced me to think about landscape ecology and the metapopulation dynamics of "lesser" organisms. Her support, financial and otherwise, was extraordinary, and a nicer person I have never met. Dr. Jon Allen exposed me to population modeling from an entomologists perspective. His enthusiasm for teaching and helping graduate students solve technical problems was exceptional. The staff at Archbold Biological Station made my stay there rewarding beyond words. Working with Dr. Reed Bowman at the Avon Park Bombing Range, and learning iii about his suburban Florida Scrub-Jays, was most stimulating. His invitation to participate in the Brevard county Habitat Conservation Plan scientific committee was most appreciated. Special thanks go to Steve Friedman and Roberta Pickert for their help with GIS problems. Dan Childs, former manager of the affiliated MacArthur Agricultural Research Center, and his staff provided unfailing assistance in many ways. Current Archbold director Dr. Hilary Swain provided generous travel support for the 1997 AOU meeting. For collecting the bulk of the demographic data used in chapter 3, 1 am indebted to Reed Bowman, Doug Stotz, Larry Riopelle, Natalie Hamel, and Mike McMillan. I thank the staff at the Natural Resources Office at the Avon Park Air Force Range, especially Bob Progulske, Paul Ebersbach, and Pat Walsh for their support. Dave McDonald was instrumental in recruiting me to work on the statewide jay survey and exposing me to the opportunities at Archbold. Dr. Keith Tarvin and Dr. Curt Atkinson kindly lent me radiotelemetry equipment and provided much valuable advice. Special thanks go to Dr. Ron Mumme who kindly allowed me to use color banded jays from the south tract of Archbold which he has been monitoring for many years. His prior work in banding, taming, and sexing these jays was critical to identifying candidates for the displacement experiment. Bill Pranty worked tirelessly with me to digitize major portions of the statewide survey. Steve Schoech provided occasional field support and many hours of entertainment on the Archbold tennis court. For stimulating discussions about jays and landscape rules, I am indebted to many colleagues engaged in the study of Florida Scrub-Jays and their conservation. Participants in the Habitat Conservation Planning group whom I have not yet mentioned include iv David Breininger, Grace Iverson, Michael O'Connell, Parks Small, Jon Thaxton, and Brian Toland. The following people contributed data for the statewide survey: Reed Bowman, Dave Breininger, Jack Dozier, Florida Game and Fresh Water Fish Commission personnel, Grace Iverson, David McDonald, Ron Mumme, Ocala National Forest personnel, Bill Pranty, Hilary Swain, Jon Thaxton, and Brian Toland. I thank David Wesley and Dawn Zattau, of the U.S. Fish and Wildlife Service, who provided lead funding for the statewide survey and helped stimulate our discussions of habitat conservation planning. For special assistance with site-specific questions related to the maps in chapter 5: Mary Barnwell, Jim Beever, Reed Bowman, Dave Breininger, Mike Eagen, Mary Huffman, Grace Iverson, Mike Jennings, Laura Lowry, Dan Pearson, Gary Popotnik, Bill Pranty, Park Smalls, Hank Smith, Jon Thaxton, Brian Toland, Jane Tutin. For assistance with GIS data: Reed Bowman, Dave Breininger, Kathy Bronson, Beth Needham, Bill Pranty, Roberta Pickert, Katy NeSmith. For advice regarding modeling: Reed Bowman, Dave Breininger, John Fitzpatrick, Glen Woolfenden. Bill Pranty deserves special thanks for the exceptionally detailed information he has collated on scrub-jay locations around the state (Pranty et al. manuscript). Input from members of the Recovery Team was most helpful. Thanks go to the U.S.F.W.S. for funding the research in chapter 5, and especially to Dawn Zattau for her support and patience. I thank the support staff at the University of Florida Department of Wildlife Ecology and Conservation. Joe Gasper provided extraordinary computer support. Leonard Pearlstein and the entire USFWS coop unit helped with innumerable computer problems. I thank support staff at Circa computing, especially Jiannong Xin for v programming resources, and John Dixon for statistical help. Dr. Ken Portier also provided valuable statistical advice. Last, but no least, I thank my family and especially my wife, Ellen Thorns. vi 1 TABLE OF CONTENTS page ACKNOWLEDGMENTS iii LIST OF TABLES xii LIST OF FIGURES xv ABSTRACT CHAPTERS 1 INTRODUCTION ! Historical Background 2 Biological Background 3 Objectives 4 2 CLASSIFYING FLORIDA SCRUB-JAY METAPOPULATIONS 9 Introduction g Statewide Survey of the Florida Scrub- Jay 1 Statewide Survey: Methods Statewide Survey: Results 13 A Method for Classifying Metapopulations 14 Metapopulation " Structure of the Florida Scrub-Jay 18 Dispersal Distances 19 Patch Occupancy 20 Population Viability Analysis 22 Metapopulation Structure 23 Caveats 25 REMOTE 3 SENSING OF FLORIDA SCRUB-JAY HABITAT 40 Introduction 4q Methods 42 Image Source 42 Image Scanning and Conversion 43 Image Rectification and Mosaicing 43 vii Image Classification 44 Manual Editing of Classification 45 Assessment of Classification Accuracy 46 Digitization of Territories and Background Features 47 Tree Cover Buffering Procedure 47 Habitat Quality Model 4g Collection of Demographic Data 49 Habitat-Demographic Analysis 49 Results " 50 Discussion 54 4 MODELING DISPERSAL IN THE FLORIDA SCRUB-JAY 86 Introduction g^ Dispersal Strategies g9 Dispersal ' Traits of the Florida and Western Scrub-Jay 90 Methods " ^ General Approach 92 GIS Files 92 Simulating Philopatric Dispersal 93 Short distance dispersal algorithm development and calibration 94 Simulating Long Distance Dispersal 95 Estimating floater mortality and mobility 96 Habitat attractiveness 9-7 Floater detection radius 97 Estimating floater frequency 9g Floater algorithm development and calibration 102 Jay displacement experiment 1 03 Constraint Analysis 1 06 Model Validation I0g Results Radiotelemetry Displacement Experiment \ 08 Floater Parameter Estimation ] 2 j Constraint Analysis 112 Calibration and Validation l\2 Discussion j j ^ 5 METAPOPULATION VIABILITY ANALYSIS OF THE FLORIDA SCRUB- JAY 132 Introduction and Objectives j 32 Methods j„ Simulation Model Description 135 Life Stages 136 Starting Population Stage Structure ZZ....... 137 " ' Annual Life Cycle 137 Territories , , A 140 viii Background Landscape Image 141 Map Production 143 Statewide metapopulation map 143 1992-1993 SMP maps ZZZZZZZZ 143 Acquisition maps ..144 GIS Database Preparation 146 Estimation ofjay populations after restoration 146 Identification of protected areas 147 Assessment of unprotected areas 147 Suburban jays 14g Simulation runs I49 Repetitions and duration of simulations 149 Reserve design configurations 150 Output statistics 151 Model Validation/Calibration 1 52 Interpreting Simulation Results 152 Results Levy (Cedar Key) (Ml) ZZZZZZZZ 155 Citrus-S.W. " Marion (M2) ZZZZZZ! 164 Pasco-Hernando (M3) 1 76 Manatee-S. Hillsborough (M4) ' 1 g6 Sarasota- W. Charlotte (M5) 196 N. W. Charlotte (M6) ZZZZZZZZZZ! 204 Central Charlotte (M7) 212 Lee and N. Collier (M8) 220 Flagler-N.E. Volusia (M9) ZZZZZZZZZZZZZZ 228 Merritt Island-S.E. Volusia and (M 1 0) 236 N. Brevard (Ml 1) ZZZZZZZ 244 Central Brevard (M 1 2) 253 S. Brevard-Indian River-N. St. Lucie (M13) ZZ! Z 261 St. Lucie -N. Martin (Ml 4) 270 Martin and N. Palm Beach (Ml 5) 278 South Palm Beach (Ml 6) ZZZZZZ ZZZZ! 286 Ocala National Forest (M 1 7) 294 N.E. Lake (Ml 8) ZZZZ 302 S.W. Volusia (Ml 9) ZZZZZZZZZ 310 Central Lake (M20) I!!!!!!!!!!!! 318 Lake Wales Ridge (M21) ZZZZZZZZZZZZZZZ" 326 Other Metapopulations 344 Brevard barrier island 344 Clay county 344 Osceola "" 344 Western Polk ^45 Bright Hour Ranch 345 Recommendations 346 Ranking Metapopulation Vulnerability 346 ix Summary of Recommendations 349 Discussion ..359 6 SYNTHESIS 365 REFERENCES 372 BIOGRAPHICAL SKETCH 383 x LIST OF TABLES MS ^ 2- 1. Summary information for 42 Florida Scrub-Jay metapopulations, including type of metapopulation, number of territories (pairs