EFFECTS OF INTRODUCED PEACOCK CICHLIDS CICHLA OCELLARIS ON NATIVE LARGEMOUTH BASS MICROPTERUS SALMOIDES IN SOUTHEAST FLORIDA By JEFFREY E. HILL 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 2003 ACKNOWLEDGMENTS Many people provided tremendous assistance in the completion of this project. All deserve my acknowledgment. I apologize beforehand to anyone inadvertently omitted. The primary acknowledgment goes to my wife, Susan, for her unfailing support throughout my entire graduate career. I thank her for sacrificing in order for me to fulfill our shared goal. 1 especially thank her for putting up with my obsession for fish. I am gratefial for the constant support of my family—^my father, mother, and sister (Baker, Jacqueline, and Kim). They shared the dream of my doctorate—I am thankfial to have fulfilled our collective aspiration. I greatly appreciate the guidance and support of my doctoral committee—Drs. Charles E. Cichra (Chair), Carter R. Gilbert, William J. Lindberg, Leo G. Nico, and Craig W. Osenberg. It was a great pleasure to work for Dr. Cichra—the experience I received in extension, research, and teaching, along with strong mentorship and his fiiendship, were instrumental in my professional development. Carter Gilbert has been a tremendous influence and is one of my real "fish heroes". I especially thank Carter for staying involved in my graduate education after his retirement—that meant a lot to me. Leo Nico's remarkable field experience with nonindigenous fishes and south Florida systems was invaluable. The many discussions (and some disagreements) over nonindigenous fishes with Leo helped shape my scientific philosophy. I appreciate the support and ii encouragement that Bill Lindberg always gave. Our discussions inside and outside of the classroom about ecology and the philosophy of science (e.g., tautology!) helped open my eyes to the wider world of science. I thank Craig Osenberg for agreeing to help out on the committee of someone studying a crazy system like the south Florida canals. I especially appreciate his assistance with preparing for my qualifying exam and his extraordinary input during the revision process. I have learned a great deal from each committee member and thank them for being a part of my graduate education. A special acknowledgment is due to Jeff Sowards for all of the hard work in the field. Thirty-six hour "days" can be rough, but he never complained. Jeff was instrumental in the field work, in the lab, and as an "ideas man" to keep the project doable. I could not have done it without his help. Also, Sharon Fitz-Coy was a tremendous help in, among other things, collecting prey for experiments and identifying insects in fish stomach contents. She was always willing to help. Special thanks are due to Paul Shafland (Florida Fish and Wildlife Conservation Commission; FWC) for a great deal of assistance, facilitation, literature, and advice during this project. Paul's largest contributions, however, came during many discussions of scientific philosophy regarding nonindigenous fish. Craig Watson and Roy Yanong (Tropical Aquaculture Laboratory) provided exceptional support throughout the process. They made the tribulations more bearable. Debbie Pouder facilitated all work done at the Sam Mitchell Aquaculture Demonstration Farm (SMADF) in Blountstown, Florida. I appreciate her help and that of the staff, especially Randall Kent. iii Ken Portier of the Department of Statistics at the University of Florida helped with the design and analysis of experiments in Chapter 5. I appreciate the hospitality of Don and Judy Bemecker who provided a great place to stay and many delicious meals during field work. Many others are due thanks for their contributions. At the University of Florida, Department of Fisheries and Aquatic Sciences, Mike Allen, Mary Cichra, Doug CoUe, Ruth Francis-Floyd, Tom Glancy, Scott Graves, Robert Leonard, Allen Riggs, Beth Sargent, Troy Thompson, Larry Tolbert, and Dan VanGenechten helped with various aspects of the project. Kelly Jacoby drew the maps. Susan Morgan greatly facilitated the graduation process. At the Florida Caribbean Science Center, U.S. Geological Survey, Gary Hill, Howard Jelks, and Bill Stranghoener assisted with facilities and equipment. Frank Morello, Jerry Krummrich, and Fred Cross (FWC) issued scientific collecting permits; Ted Hoehn (FWC) kindly provided access and instruction to the FWC fish database; Eric Nagid (FWC) helped answer questions about use and content of the database. I gratefully acknowledge the financial and facilities assistance of the Department of Fisheries and Aquatic Sciences, the Sam Mitchell Aquaculture Demonstration Farm, and the Tropical Aquaculture Laboratory of the University of Florida. Financial assistance was further provided by a Dean's Fellowship for Graduate Research from the Institute of Food and Agricultural Sciences, University of Florida, the Florida Department of Agriculture and Consumer Services, by Ryan Kelley Memorial Scholarships from the International Women's Fishing Association, and the Roger Rottmann Memorial Scholarship from the Florida Chapter, American Fisheries Society. iv TABLE OF CONTENTS gage ACKNOWLEDGMENTS ii ABSTRACT vii CHAPTER 1 INTRODUCTION 1 2 STUDY SYSTEM 11 Study Species 11 Canal System 14 Study Sites 18 3 A QUALITATIVE RISK ASSESSMENT OF PEACOCK CICHLIDS CICHLA OCELLARIS AND SOME IMPLICATIONS FOR THE METHODOLOGY 24 Introduction 24 Background Information 27 Qualitative Risk Assessment 36 Discussion 43 4 ONTOGENETIC DIETARY SHIFTS AND DIETARY OVERLAP OF A NONINDIGENOUS FISH AND AN ECOMORPHOLOGICALLY SIMILAR NATIVE FISH 49 Introduction 49 Methods 51 Results 56 Discussion 68 5 EXPERIMENTAL PREY HANDLING AND SELECTIVITY OF TWO MORPHOLOGICALLY SIMILAR PREDATORY FISHES- NATIVE LARGEMOUTH BASS AND INTRODUCED PEACOCK CICHLIDS 79 V Introduction 79 Methods 82 Results 90 Discussion 99 6 THE INTRODUCTION OF PEACOCK CICHLIDS INTO SOUTHEAST FLORIDA: POPULATION-LEVEL CONSEQUENCES FOR LARGEMOUTH BASS 109 Introduction 1 09 Methods 112 Results 114 Discussion 119 7 SUMMARY 127 REFERENCES 133 BIOGRAPHICAL SKETCH 154 vi Abstract of Dissertation Presented to the Graduate School of the University of Florida in Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy EFFECTS OF INTRODUCED PEACOCK CICHLIDS CICHLA OCELLARIS ON NATIVE LARGEMOUTH BASS MICROPTERUS SALMOIDES IN SOUTHEAST FLORIDA By Jeffrey E. Hill December 2003 Chair: Charles E. Cichra Major Department: Fisheries and Aquatic Sciences The predatory peacock cichlid Cichla ocellaris , native to South America, was introduced into southeast Florida to enhance recreational fishing and to provide biological control over nonindigenous fishes. This action was controversial and there is concern over the effects of the introduction on native species, especially the ecomorphologically similar largemouth bass Micropterus salmoides . I conducted a formal qualitative risk assessment to evaluate the risks associated with the peacock cichlid introducfion and to test predictions of the algorithms. The esfimate of risk for peacock cichlids was Low. The evaluation elucidated shortcomings in the methodology (e.g., emphasis on a single output) and I made recommendations for its improvement. Part of the concern over the peacock cichlid introduction is their similarity to largemouth bass. I tested and found support for the hypothesis that morphological similarity equals similarity in prey handling ability and prey selection. I documented substantial overlap in vii trophic morphology and dietary ontogeny between juveniles and found similar prey handling ability and prey selection by adults in tank experiments. These results suggest that largemouth bass and peacock cichlids have similar effects on the prey base. However, largemouth bass had exclusive use of highly defensive prey (i.e., crayfish). Largemouth bass populations might be expected to have declined, given evidence of direct negative effects of peacock cichlids. I investigated this question by comparing largemouth bass populations in southeast Florida canals with populations in lakes and streams (statewide and in south Florida). Additionally, I compared canals with peacock cichlids to canals without peacock cichlids. The data did not support a hypothesis of declines in largemouth bass abundance due to peacock cichlids. These results question common assumptions concerning the effects of nonindigenous species— 1) that direct effects of predation and competition are the predominant effects and 2) the narrow view that invaded communities are organismic (i.e., Clementsian)—and challenge the uncritical acceptance of the premise that nonindigenous fishes will invariably cause ecological harm. My results suggest that other factors (e.g., indirect effects) may be important and that invasion biology should develop a pluralistic theory that incorporates Gleasonian concepts (i.e., individualistic communities). viii CHAPTER 1 INTRODUCTION Introduced species are considered threats to natural ecosystems and biodiversity worldwide (Lodge 1993). For example, Miller et al. (1989) reported that nonindigenous species were a factor in the extinction of 27 North American fishes (68% of extinctions). Similarly, Lassuy (1995) stated that nonindigenous species were factors in listing 48 imperiled freshwater fishes under the Endangered Species Act in the USA (70% of listed fishes). Although ecological